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• 42.1: The Nervous System Organization
Nervous systems throughout the animal kingdom vary in structure and complexity, as illustrated by the variety of animals shown in Figure 35.1.1. Some organisms, like sea sponges, lack a true nervous system. Others, like jellyfish, lack a true brain and instead have a system of separate but connected nerve cells (neurons) called a “nerve net.” Echinoderms such as sea stars have nerve cells that are bundled into fibers called nerves.
• 42.2: The Mechanism of Nerve Impulse Transmission
All functions performed by the nervous system—from a simple motor reflex to more advanced functions like making a memory or a decision—require neurons to communicate with one another. While humans use words and body language to communicate, neurons use electrical and chemical signals. Just like a person in a committee, one neuron usually receives and synthesizes messages from multiple other neurons before “making the decision” to send the message on to other neurons.
• 42.3: Synapses- Where Neurons Communicate With Other Cells
The central nervous system is made up of the brain and spinal cord and is covered with three layers of protective coverings called meninges (from the Greek word for membrane). The outermost layer is the dura mater with the primary function for this thick layer is to protect the brain and spinal cord. The dura mater also contains vein-like structures that carry blood from the brain back to the heart. The middle layer is the web-like arachnoid mater. The last layer is the pia mater.
• 42.4: The Central Nervous System- Brain and Spinal Cord
The central nervous system is made up of the brain and spinal cord and is covered with three layers of protective coverings called meninges (from the Greek word for membrane). The outermost layer is the dura mater with the primary function for this thick layer is to protect the brain and spinal cord. The dura mater also contains vein-like structures that carry blood from the brain back to the heart. The middle layer is the web-like arachnoid mater. The last layer is the pia mater.
• 42.5: The Peripheral Nervous System- Spinal and Cranial Nerves
The peripheral nervous system (PNS) is the connection between the central nervous system and the rest of the body. The central nervious system (CNS) is like the power plant of the nervous system. It creates the signals that control the functions of the body. The PNS is like the wires that go to individual houses. Without those “wires,” the signals produced by the CNS could not control the body (and the CNS would not be able to receive sensory information from the body either).
42: The Nervous System
Skills to Develop
• List and describe the functions of the structural components of a neuron
• List and describe the four main types of neurons
• Compare the functions of different types of glial cells
Nervous systems throughout the animal kingdom vary in structure and complexity, as illustrated by the variety of animals shown in Figure \(1\). Some organisms, like sea sponges, lack a true nervous system. Others, like jellyfish, lack a true brain and instead have a system of separate but connected nerve cells (neurons) called a “nerve net.” Echinoderms such as sea stars have nerve cells that are bundled into fibers called nerves. Flatworms of the phylum Platyhelminthes have both a central nervous system (CNS), made up of a small “brain” and two nerve cords, and a peripheral nervous system (PNS) containing a system of nerves that extend throughout the body. The insect nervous system is more complex but also fairly decentralized. It contains a brain, ventral nerve cord, and ganglia (clusters of connected neurons). These ganglia can control movements and behaviors without input from the brain. Octopi may have the most complicated of invertebrate nervous systems—they have neurons that are organized in specialized lobes and eyes that are structurally similar to vertebrate species.
Compared to invertebrates, vertebrate nervous systems are more complex, centralized, and specialized. While there is great diversity among different vertebrate nervous systems, they all share a basic structure: a CNS that contains a brain and spinal cord and a PNS made up of peripheral sensory and motor nerves. One interesting difference between the nervous systems of invertebrates and vertebrates is that the nerve cords of many invertebrates are located ventrally whereas the vertebrate spinal cords are located dorsally. There is debate among evolutionary biologists as to whether these different nervous system plans evolved separately or whether the invertebrate body plan arrangement somehow “flipped” during the evolution of vertebrates.
Link to Learning
Watch this video of biologist Mark Kirschner discussing the “flipping” phenomenon of vertebrate evolution.
The nervous system is made up of neurons, specialized cells that can receive and transmit chemical or electrical signals, and glia, cells that provide support functions for the neurons by playing an information processing role that is complementary to neurons. A neuron can be compared to an electrical wire—it transmits a signal from one place to another. Glia can be compared to the workers at the electric company who make sure wires go to the right places, maintain the wires, and take down wires that are broken. Although glia have been compared to workers, recent evidence suggests that also usurp some of the signaling functions of neurons.
There is great diversity in the types of neurons and glia that are present in different parts of the nervous system. There are four major types of neurons, and they share several important cellular components.
Neurons
The nervous system of the common laboratory fly, Drosophila melanogaster, contains around 100,000 neurons, the same number as a lobster. This number compares to 75 million in the mouse and 300 million in the octopus. A human brain contains around 86 billion neurons. Despite these very different numbers, the nervous systems of these animals control many of the same behaviors—from basic reflexes to more complicated behaviors like finding food and courting mates. The ability of neurons to communicate with each other as well as with other types of cells underlies all of these behaviors.
Most neurons share the same cellular components. But neurons are also highly specialized—different types of neurons have different sizes and shapes that relate to their functional roles.
Parts of a Neuron
Like other cells, each neuron has a cell body (or soma) that contains a nucleus, smooth and rough endoplasmic reticulum, Golgi apparatus, mitochondria, and other cellular components. Neurons also contain unique structures, illustrated in Figure \(2\) for receiving and sending the electrical signals that make neuronal communication possible. Dendrites are tree-like structures that extend away from the cell body to receive messages from other neurons at specialized junctions called synapses. Although some neurons do not have any dendrites, some types of neurons have multiple dendrites. Dendrites can have small protrusions called dendritic spines, which further increase surface area for possible synaptic connections.
Once a signal is received by the dendrite, it then travels passively to the cell body. The cell body contains a specialized structure, the axon hillock that integrates signals from multiple synapses and serves as a junction between the cell body and an axon. An axon is a tube-like structure that propagates the integrated signal to specialized endings called axon terminals. These terminals in turn synapse on other neurons, muscle, or target organs. Chemicals released at axon terminals allow signals to be communicated to these other cells. Neurons usually have one or two axons, but some neurons, like amacrine cells in the retina, do not contain any axons. Some axons are covered with myelin, which acts as an insulator to minimize dissipation of the electrical signal as it travels down the axon, greatly increasing the speed on conduction. This insulation is important as the axon from a human motor neuron can be as long as a meter—from the base of the spine to the toes. The myelin sheath is not actually part of the neuron. Myelin is produced by glial cells. Along the axon there are periodic gaps in the myelin sheath. These gaps are called nodes of Ranvier and are sites where the signal is “recharged” as it travels along the axon.
It is important to note that a single neuron does not act alone—neuronal communication depends on the connections that neurons make with one another (as well as with other cells, like muscle cells). Dendrites from a single neuron may receive synaptic contact from many other neurons. For example, dendrites from a Purkinje cell in the cerebellum are thought to receive contact from as many as 200,000 other neurons.
Art Connection
Which of the following statements is false?
1. The soma is the cell body of a nerve cell.
2. Myelin sheath provides an insulating layer to the dendrites.
3. Axons carry the signal from the soma to the target.
4. Dendrites carry the signal to the soma.
Types of Neurons
There are different types of neurons, and the functional role of a given neuron is intimately dependent on its structure. There is an amazing diversity of neuron shapes and sizes found in different parts of the nervous system (and across species), as illustrated by the neurons shown in Figure \(3\).
While there are many defined neuron cell subtypes, neurons are broadly divided into four basic types: unipolar, bipolar, multipolar, and pseudounipolar. Figure \(4\) illustrates these four basic neuron types. Unipolar neurons have only one structure that extends away from the soma. These neurons are not found in vertebrates but are found in insects where they stimulate muscles or glands. A bipolar neuron has one axon and one dendrite extending from the soma. An example of a bipolar neuron is a retinal bipolar cell, which receives signals from photoreceptor cells that are sensitive to light and transmits these signals to ganglion cells that carry the signal to the brain. Multipolar neurons are the most common type of neuron. Each multipolar neuron contains one axon and multiple dendrites. Multipolar neurons can be found in the central nervous system (brain and spinal cord). An example of a multipolar neuron is a Purkinje cell in the cerebellum, which has many branching dendrites but only one axon. Pseudounipolar cells share characteristics with both unipolar and bipolar cells. A pseudounipolar cell has a single process that extends from the soma, like a unipolar cell, but this process later branches into two distinct structures, like a bipolar cell. Most sensory neurons are pseudounipolar and have an axon that branches into two extensions: one connected to dendrites that receive sensory information and another that transmits this information to the spinal cord.
Everyday Connection: Neurogenesis
At one time, scientists believed that people were born with all the neurons they would ever have. Research performed during the last few decades indicates that neurogenesis, the birth of new neurons, continues into adulthood. Neurogenesis was first discovered in songbirds that produce new neurons while learning songs. For mammals, new neurons also play an important role in learning: about 1000 new neurons develop in the hippocampus (a brain structure involved in learning and memory) each day. While most of the new neurons will die, researchers found that an increase in the number of surviving new neurons in the hippocampus correlated with how well rats learned a new task. Interestingly, both exercise and some antidepressant medications also promote neurogenesis in the hippocampus. Stress has the opposite effect. While neurogenesis is quite limited compared to regeneration in other tissues, research in this area may lead to new treatments for disorders such as Alzheimer’s, stroke, and epilepsy.
How do scientists identify new neurons? A researcher can inject a compound called bromodeoxyuridine (BrdU) into the brain of an animal. While all cells will be exposed to BrdU, BrdU will only be incorporated into the DNA of newly generated cells that are in S phase. A technique called immunohistochemistry can be used to attach a fluorescent label to the incorporated BrdU, and a researcher can use fluorescent microscopy to visualize the presence of BrdU, and thus new neurons, in brain tissue. Figure \(5\) is a micrograph which shows fluorescently labeled neurons in the hippocampus of a rat.
Link to Learning
This site contains more information about neurogenesis, including an interactive laboratory simulation and a video that explains how BrdU labels new cells.
Glia
While glia are often thought of as the supporting cast of the nervous system, the number of glial cells in the brain actually outnumbers the number of neurons by a factor of ten. Neurons would be unable to function without the vital roles that are fulfilled by these glial cells. Glia guide developing neurons to their destinations, buffer ions and chemicals that would otherwise harm neurons, and provide myelin sheaths around axons. Scientists have recently discovered that they also play a role in responding to nerve activity and modulating communication between nerve cells. When glia do not function properly, the result can be disastrous—most brain tumors are caused by mutations in glia.
Types of Glia
There are several different types of glia with different functions, two of which are shown in Figure \(6\). Astrocytes, shown in Figure \(7\) make contact with both capillaries and neurons in the CNS. They provide nutrients and other substances to neurons, regulate the concentrations of ions and chemicals in the extracellular fluid, and provide structural support for synapses. Astrocytes also form the blood-brain barrier—a structure that blocks entrance of toxic substances into the brain. Astrocytes, in particular, have been shown through calcium imaging experiments to become active in response to nerve activity, transmit calcium waves between astrocytes, and modulate the activity of surrounding synapses.
Satellite glia provide nutrients and structural support for neurons in the PNS. Microglia scavenge and degrade dead cells and protect the brain from invading microorganisms. Oligodendrocytes, shown in Figure \(7\) form myelin sheaths around axons in the CNS. One axon can be myelinated by several oligodendrocytes, and one oligodendrocyte can provide myelin for multiple neurons. This is distinctive from the PNS where a single Schwann cell provides myelin for only one axon as the entire Schwann cell surrounds the axon. Radial glia serve as scaffolds for developing neurons as they migrate to their end destinations. Ependymal cells line fluid-filled ventricles of the brain and the central canal of the spinal cord. They are involved in the production of cerebrospinal fluid, which serves as a cushion for the brain, moves the fluid between the spinal cord and the brain, and is a component for the choroid plexus.
Summary
The nervous system is made up of neurons and glia. Neurons are specialized cells that are capable of sending electrical as well as chemical signals. Most neurons contain dendrites, which receive these signals, and axons that send signals to other neurons or tissues. There are four main types of neurons: unipolar, bipolar, multipolar, and pseudounipolar neurons. Glia are non-neuronal cells in the nervous system that support neuronal development and signaling. There are several types of glia that serve different functions.
Art Connections
Figure \(2\): Which of the following statements is false?
1. The soma is the cell body of a nerve cell.
2. Myelin sheath provides an insulating layer to the dendrites.
3. Axons carry the signal from the soma to the target.
4. Dendrites carry the signal to the soma.
Answer
B
Glossary
astrocyte
glial cell in the central nervous system that provide nutrients, extracellular buffering, and structural support for neurons; also makes up the blood-brain barrier
axon
tube-like structure that propagates a signal from a neuron’s cell body to axon terminals
axon hillock
electrically sensitive structure on the cell body of a neuron that integrates signals from multiple neuronal connections
axon terminal
structure on the end of an axon that can form a synapse with another neuron
dendrite
structure that extends away from the cell body to receive messages from other neurons
ependymal
cell that lines fluid-filled ventricles of the brain and the central canal of the spinal cord; involved in production of cerebrospinal fluid
glia
(also, glial cells) cells that provide support functions for neurons
microglia
glia that scavenge and degrade dead cells and protect the brain from invading microorganisms
myelin
fatty substance produced by glia that insulates axons
neuron
specialized cell that can receive and transmit electrical and chemical signals
nodes of Ranvier
gaps in the myelin sheath where the signal is recharged
oligodendrocyte
glial cell that myelinates central nervous system neuron axons
radial glia
glia that serve as scaffolds for developing neurons as they migrate to their final destinations
satellite glia
glial cell that provides nutrients and structural support for neurons in the peripheral nervous system
Schwann cell
glial cell that creates myelin sheath around a peripheral nervous system neuron axon
synapse
junction between two neurons where neuronal signals are communicated
42.01: The Nervous System Organization
Skills to Develop
• Describe the organization and functions of the sympathetic and parasympathetic nervous systems
• Describe the organization and function of the sensory-somatic nervous system
The peripheral nervous system (PNS) is the connection between the central nervous system and the rest of the body. The CNS is like the power plant of the nervous system. It creates the signals that control the functions of the body. The PNS is like the wires that go to individual houses. Without those “wires,” the signals produced by the CNS could not control the body (and the CNS would not be able to receive sensory information from the body either).
The PNS can be broken down into the autonomic nervous system, which controls bodily functions without conscious control, and the sensory-somatic nervous system, which transmits sensory information from the skin, muscles, and sensory organs to the CNS and sends motor commands from the CNS to the muscles.
Autonomic Nervous System
Art Connection
Which of the following statements is false?
1. The parasympathetic pathway is responsible for resting the body, while the sympathetic pathway is responsible for preparing for an emergency.
2. Most preganglionic neurons in the sympathetic pathway originate in the spinal cord.
3. Slowing of the heartbeat is a parasympathetic response.
4. Parasympathetic neurons are responsible for releasing norepinephrine on the target organ, while sympathetic neurons are responsible for releasing acetylcholine.
The autonomic nervous system serves as the relay between the CNS and the internal organs. It controls the lungs, the heart, smooth muscle, and exocrine and endocrine glands. The autonomic nervous system controls these organs largely without conscious control; it can continuously monitor the conditions of these different systems and implement changes as needed. Signaling to the target tissue usually involves two synapses: a preganglionic neuron (originating in the CNS) synapses to a neuron in a ganglion that, in turn, synapses on the target organ, as illustrated in Figure \(1\). There are two divisions of the autonomic nervous system that often have opposing effects: the sympathetic nervous system and the parasympathetic nervous system.
Sympathetic Nervous System
The sympathetic nervous system is responsible for the “fight or flight” response that occurs when an animal encounters a dangerous situation. One way to remember this is to think of the surprise a person feels when encountering a snake (“snake” and “sympathetic” both begin with “s”). Examples of functions controlled by the sympathetic nervous system include an accelerated heart rate and inhibited digestion. These functions help prepare an organism’s body for the physical strain required to escape a potentially dangerous situation or to fend off a predator.
Most preganglionic neurons in the sympathetic nervous system originate in the spinal cord, as illustrated in Figure \(2\). The axons of these neurons release acetylcholine on postganglionic neurons within sympathetic ganglia (the sympathetic ganglia form a chain that extends alongside the spinal cord). The acetylcholine activates the postganglionic neurons. Postganglionic neurons then release norepinephrine onto target organs. As anyone who has ever felt a rush before a big test, speech, or athletic event can attest, the effects of the sympathetic nervous system are quite pervasive. This is both because one preganglionic neuron synapses on multiple postganglionic neurons, amplifying the effect of the original synapse, and because the adrenal gland also releases norepinephrine (and the closely related hormone epinephrine) into the blood stream. The physiological effects of this norepinephrine release include dilating the trachea and bronchi (making it easier for the animal to breathe), increasing heart rate, and moving blood from the skin to the heart, muscles, and brain (so the animal can think and run). The strength and speed of the sympathetic response helps an organism avoid danger, and scientists have found evidence that it may also increase LTP—allowing the animal to remember the dangerous situation and avoid it in the future.
Parasympathetic Nervous System
While the sympathetic nervous system is activated in stressful situations, the parasympathetic nervous system allows an animal to “rest and digest.” One way to remember this is to think that during a restful situation like a picnic, the parasympathetic nervous system is in control (“picnic” and “parasympathetic” both start with “p”). Parasympathetic preganglionic neurons have cell bodies located in the brainstem and in the sacral (toward the bottom) spinal cord, as shown in Figure \(2\). The axons of the preganglionic neurons release acetylcholine on the postganglionic neurons, which are generally located very near the target organs. Most postganglionic neurons release acetylcholine onto target organs, although some release nitric oxide.
The parasympathetic nervous system resets organ function after the sympathetic nervous system is activated (the common adrenaline dump you feel after a ‘fight-or-flight’ event). Effects of acetylcholine release on target organs include slowing of heart rate, lowered blood pressure, and stimulation of digestion.
Sensory-Somatic Nervous System
The sensory-somatic nervous system is made up of cranial and spinal nerves and contains both sensory and motor neurons. Sensory neurons transmit sensory information from the skin, skeletal muscle, and sensory organs to the CNS. Motor neurons transmit messages about desired movement from the CNS to the muscles to make them contract. Without its sensory-somatic nervous system, an animal would be unable to process any information about its environment (what it sees, feels, hears, and so on) and could not control motor movements. Unlike the autonomic nervous system, which has two synapses between the CNS and the target organ, sensory and motor neurons have only one synapse—one ending of the neuron is at the organ and the other directly contacts a CNS neuron. Acetylcholine is the main neurotransmitter released at these synapses.
Humans have 12 cranial nerves, nerves that emerge from or enter the skull (cranium), as opposed to the spinal nerves, which emerge from the vertebral column. Each cranial nerve is accorded a name, which are detailed in Figure \(3\). Some cranial nerves transmit only sensory information. For example, the olfactory nerve transmits information about smells from the nose to the brainstem. Other cranial nerves transmit almost solely motor information. For example, the oculomotor nerve controls the opening and closing of the eyelid and some eye movements. Other cranial nerves contain a mix of sensory and motor fibers. For example, the glossopharyngeal nerve has a role in both taste (sensory) and swallowing (motor).
Spinal nerves transmit sensory and motor information between the spinal cord and the rest of the body. Each of the 31 spinal nerves (in humans) contains both sensory and motor axons. The sensory neuron cell bodies are grouped in structures called dorsal root ganglia and are shown in Figure \(4\). Each sensory neuron has one projection—with a sensory receptor ending in skin, muscle, or sensory organs—and another that synapses with a neuron in the dorsal spinal cord. Motor neurons have cell bodies in the ventral gray matter of the spinal cord that project to muscle through the ventral root. These neurons are usually stimulated by interneurons within the spinal cord but are sometimes directly stimulated by sensory neurons.
Summary
The peripheral nervous system contains both the autonomic and sensory-somatic nervous systems. The autonomic nervous system provides unconscious control over visceral functions and has two divisions: the sympathetic and parasympathetic nervous systems. The sympathetic nervous system is activated in stressful situations to prepare the animal for a “fight or flight” response. The parasympathetic nervous system is active during restful periods. The sensory-somatic nervous system is made of cranial and spinal nerves that transmit sensory information from skin and muscle to the CNS and motor commands from the CNS to the muscles.
Art Connections
Figure \(1\): Which of the following statements is false?
1. The parasympathetic pathway is responsible for relaxing the body, while the sympathetic pathway is responsible for preparing for an emergency.
2. Most preganglionic neurons in the sympathetic pathway originate in the spinal cord.
3. Slowing of the heartbeat is a parasympathetic response.
4. Parasympathetic neurons are responsible for releasing norepinephrine on the target organ, while sympathetic neurons are responsible for releasing acetylcholine.
Answer
D
Glossary
acetylcholine
neurotransmitter released by neurons in the central nervous system and peripheral nervous system
autonomic nervous system
part of the peripheral nervous system that controls bodily functions
cranial nerve
sensory and/or motor nerve that emanates from the brain
norepinephrine
neurotransmitter and hormone released by activation of the sympathetic nervous system
parasympathetic nervous system
division of autonomic nervous system that regulates visceral functions during rest and digestion
sensory-somatic nervous system
system of sensory and motor nerves
spinal nerve
nerve projecting between skin or muscle and spinal cord
sympathetic nervous system
division of autonomic nervous system activated during stressful “fight or flight” situations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/42%3A_The_Nervous_System/42.01%3A_The_Nervous_System_Organization/42.1.01%3A_The_Peripheral_Nervous_System.txt |
Skills to Develop
• Describe the basis of the resting membrane potential
• Explain the stages of an action potential and how action potentials are propagated
• Explain the similarities and differences between chemical and electrical synapses
• Describe long-term potentiation and long-term depression
All functions performed by the nervous system—from a simple motor reflex to more advanced functions like making a memory or a decision—require neurons to communicate with one another. While humans use words and body language to communicate, neurons use electrical and chemical signals. Just like a person in a committee, one neuron usually receives and synthesizes messages from multiple other neurons before “making the decision” to send the message on to other neurons.
Nerve Impulse Transmission within a Neuron
For the nervous system to function, neurons must be able to send and receive signals. These signals are possible because each neuron has a charged cellular membrane (a voltage difference between the inside and the outside), and the charge of this membrane can change in response to neurotransmitter molecules released from other neurons and environmental stimuli. To understand how neurons communicate, one must first understand the basis of the baseline or ‘resting’ membrane charge.
Neuronal Charged Membranes
The lipid bilayer membrane that surrounds a neuron is impermeable to charged molecules or ions. To enter or exit the neuron, ions must pass through special proteins called ion channels that span the membrane. Ion channels have different configurations: open, closed, and inactive, as illustrated in Figure \(1\). Some ion channels need to be activated in order to open and allow ions to pass into or out of the cell. These ion channels are sensitive to the environment and can change their shape accordingly. Ion channels that change their structure in response to voltage changes are called voltage-gated ion channels. Voltage-gated ion channels regulate the relative concentrations of different ions inside and outside the cell. The difference in total charge between the inside and outside of the cell is called the membrane potential.
Link to Learning
This video discusses the basis of the resting membrane potential.
Resting Membrane Potential
A neuron at rest is negatively charged: the inside of a cell is approximately 70 millivolts more negative than the outside (−70 mV, note that this number varies by neuron type and by species). This voltage is called the resting membrane potential; it is caused by differences in the concentrations of ions inside and outside the cell. If the membrane were equally permeable to all ions, each type of ion would flow across the membrane and the system would reach equilibrium. Because ions cannot simply cross the membrane at will, there are different concentrations of several ions inside and outside the cell, as shown in the table below. The difference in the number of positively charged potassium ions (K+) inside and outside the cell dominates the resting membrane potential (Figure \(2\)). When the membrane is at rest, K+ ions accumulate inside the cell due to a net movement with the concentration gradient. The negative resting membrane potential is created and maintained by increasing the concentration of cations outside the cell (in the extracellular fluid) relative to inside the cell (in the cytoplasm). The negative charge within the cell is created by the cell membrane being more permeable to potassium ion movement than sodium ion movement. In neurons, potassium ions are maintained at high concentrations within the cell while sodium ions are maintained at high concentrations outside of the cell. The cell possesses potassium and sodium leakage channels that allow the two cations to diffuse down their concentration gradient. However, the neurons have far more potassium leakage channels than sodium leakage channels. Therefore, potassium diffuses out of the cell at a much faster rate than sodium leaks in. Because more cations are leaving the cell than are entering, this causes the interior of the cell to be negatively charged relative to the outside of the cell. The actions of the sodium potassium pump help to maintain the resting potential, once established. Recall that sodium potassium pumps brings two K+ ions into the cell while removing three Na+ ions per ATP consumed. As more cations are expelled from the cell than taken in, the inside of the cell remains negatively charged relative to the extracellular fluid. It should be noted that calcium ions (Cl) tend to accumulate outside of the cell because they are repelled by negatively-charged proteins within the cytoplasm.
Table \(1\): Ion Concentration Inside and Outside Neurons. The resting membrane potential is a result of different concentrations inside and outside the cell.
Ion Extracellular concentration (mM) Intracellular concentration (mM) Ratio outside/inside
Na+ 145 12 12
K+ 4 155 0.026
Cl 120 4 30
Organic anions (A−) 100
Action Potential
A neuron can receive input from other neurons and, if this input is strong enough, send the signal to downstream neurons. Transmission of a signal between neurons is generally carried by a chemical called a neurotransmitter. Transmission of a signal within a neuron (from dendrite to axon terminal) is carried by a brief reversal of the resting membrane potential called an action potential. When neurotransmitter molecules bind to receptors located on a neuron’s dendrites, ion channels open. At excitatory synapses, this opening allows positive ions to enter the neuron and results in depolarization of the membrane—a decrease in the difference in voltage between the inside and outside of the neuron. A stimulus from a sensory cell or another neuron depolarizes the target neuron to its threshold potential (-55 mV). Na+ channels in the axon hillock open, allowing positive ions to enter the cell (Figure \(3\) and Figure \(4\)). Once the sodium channels open, the neuron completely depolarizes to a membrane potential of about +40 mV. Action potentials are considered an "all-or nothing" event, in that, once the threshold potential is reached, the neuron always completely depolarizes. Once depolarization is complete, the cell must now "reset" its membrane voltage back to the resting potential. To accomplish this, the Na+ channels close and cannot be opened. This begins the neuron's refractory period, in which it cannot produce another action potential because its sodium channels will not open. At the same time, voltage-gated K+ channels open, allowing K+ to leave the cell. As K+ ions leave the cell, the membrane potential once again becomes negative. The diffusion of K+ out of the cell actually hyperpolarizes the cell, in that the membrane potential becomes more negative than the cell's normal resting potential. At this point, the sodium channels will return to their resting state, meaning they are ready to open again if the membrane potential again exceeds the threshold potential. Eventually the extra K+ ions diffuse out of the cell through the potassium leakage channels, bringing the cell from its hyperpolarized state, back to its resting membrane potential.
Art Connection
Potassium channel blockers, such as amiodarone and procainamide, which are used to treat abnormal electrical activity in the heart, called cardiac dysrhythmia, impede the movement of K+ through voltage-gated K+ channels. Which part of the action potential would you expect potassium channels to affect?
Link to Learning
This video presents an overview of action potential.
Myelin and the Propagation of the Action Potential
For an action potential to communicate information to another neuron, it must travel along the axon and reach the axon terminals where it can initiate neurotransmitter release. The speed of conduction of an action potential along an axon is influenced by both the diameter of the axon and the axon’s resistance to current leak. Myelin acts as an insulator that prevents current from leaving the axon; this increases the speed of action potential conduction. In demyelinating diseases like multiple sclerosis, action potential conduction slows because current leaks from previously insulated axon areas. The nodes of Ranvier, illustrated in Figure \(5\) are gaps in the myelin sheath along the axon. These unmyelinated spaces are about one micrometer long and contain voltage gated Na+ and K+ channels. Flow of ions through these channels, particularly the Na+ channels, regenerates the action potential over and over again along the axon. This ‘jumping’ of the action potential from one node to the next is called saltatory conduction. If nodes of Ranvier were not present along an axon, the action potential would propagate very slowly since Na+ and K+ channels would have to continuously regenerate action potentials at every point along the axon instead of at specific points. Nodes of Ranvier also save energy for the neuron since the channels only need to be present at the nodes and not along the entire axon.
Synaptic Transmission
The synapse or “gap” is the place where information is transmitted from one neuron to another. Synapses usually form between axon terminals and dendritic spines, but this is not universally true. There are also axon-to-axon, dendrite-to-dendrite, and axon-to-cell body synapses. The neuron transmitting the signal is called the presynaptic neuron, and the neuron receiving the signal is called the postsynaptic neuron. Note that these designations are relative to a particular synapse—most neurons are both presynaptic and postsynaptic. There are two types of synapses: chemical and electrical.
Chemical Synapse
When an action potential reaches the axon terminal it depolarizes the membrane and opens voltage-gated Na+ channels. Na+ ions enter the cell, further depolarizing the presynaptic membrane. This depolarization causes voltage-gated Ca2+ channels to open. Calcium ions entering the cell initiate a signaling cascade that causes small membrane-bound vesicles, called synaptic vesicles, containing neurotransmitter molecules to fuse with the presynaptic membrane. Synaptic vesicles are shown in Figure \(6\), which is an image from a scanning electron microscope.
Fusion of a vesicle with the presynaptic membrane causes neurotransmitter to be released into the synaptic cleft, the extracellular space between the presynaptic and postsynaptic membranes, as illustrated in Figure \(7\). The neurotransmitter diffuses across the synaptic cleft and binds to receptor proteins on the postsynaptic membrane.
The binding of a specific neurotransmitter causes particular ion channels, in this case ligand-gated channels, on the postsynaptic membrane to open. Neurotransmitters can either have excitatory or inhibitory effects on the postsynaptic membrane, as detailed in the table below. For example, when acetylcholine is released at the synapse between a nerve and muscle (called the neuromuscular junction) by a presynaptic neuron, it causes postsynaptic Na+ channels to open. Na+ enters the postsynaptic cell and causes the postsynaptic membrane to depolarize. This depolarization is called an excitatory postsynaptic potential (EPSP) and makes the postsynaptic neuron more likely to fire an action potential. Release of neurotransmitter at inhibitory synapses causes inhibitory postsynaptic potentials (IPSPs), a hyperpolarization of the presynaptic membrane. For example, when the neurotransmitter GABA (gamma-aminobutyric acid) is released from a presynaptic neuron, it binds to and opens Cl- channels. Cl- ions enter the cell and hyperpolarizes the membrane, making the neuron less likely to fire an action potential.
Once neurotransmission has occurred, the neurotransmitter must be removed from the synaptic cleft so the postsynaptic membrane can “reset” and be ready to receive another signal. This can be accomplished in three ways: the neurotransmitter can diffuse away from the synaptic cleft, it can be degraded by enzymes in the synaptic cleft, or it can be recycled (sometimes called reuptake) by the presynaptic neuron. Several drugs act at this step of neurotransmission. For example, some drugs that are given to Alzheimer’s patients work by inhibiting acetylcholinesterase, the enzyme that degrades acetylcholine. This inhibition of the enzyme essentially increases neurotransmission at synapses that release acetylcholine. Once released, the acetylcholine stays in the cleft and can continually bind and unbind to postsynaptic receptors.
Table \(2\): Neurotransmitter Function and Location
Neurotransmitter Example Location
Acetylcholine CNS and/or PNS
Biogenic amine Dopamine, serotonin, norepinephrine CNS and/or PNS
Amino acid Glycine, glutamate, aspartate, gamma aminobutyric acid CNS
Neuropeptide Substance P, endorphins CNS and/or PNS
Electrical Synapse
While electrical synapses are fewer in number than chemical synapses, they are found in all nervous systems and play important and unique roles. The mode of neurotransmission in electrical synapses is quite different from that in chemical synapses. In an electrical synapse, the presynaptic and postsynaptic membranes are very close together and are actually physically connected by channel proteins forming gap junctions. Gap junctions allow current to pass directly from one cell to the next. In addition to the ions that carry this current, other molecules, such as ATP, can diffuse through the large gap junction pores.
There are key differences between chemical and electrical synapses. Because chemical synapses depend on the release of neurotransmitter molecules from synaptic vesicles to pass on their signal, there is an approximately one millisecond delay between when the axon potential reaches the presynaptic terminal and when the neurotransmitter leads to opening of postsynaptic ion channels. Additionally, this signaling is unidirectional. Signaling in electrical synapses, in contrast, is virtually instantaneous (which is important for synapses involved in key reflexes), and some electrical synapses are bidirectional. Electrical synapses are also more reliable as they are less likely to be blocked, and they are important for synchronizing the electrical activity of a group of neurons. For example, electrical synapses in the thalamus are thought to regulate slow-wave sleep, and disruption of these synapses can cause seizures.
Signal Summation
Sometimes a single EPSP is strong enough to induce an action potential in the postsynaptic neuron, but often multiple presynaptic inputs must create EPSPs around the same time for the postsynaptic neuron to be sufficiently depolarized to fire an action potential. This process is called summation and occurs at the axon hillock, as illustrated in Figure \(8\). Additionally, one neuron often has inputs from many presynaptic neurons—some excitatory and some inhibitory—so IPSPs can cancel out EPSPs and vice versa. It is the net change in postsynaptic membrane voltage that determines whether the postsynaptic cell has reached its threshold of excitation needed to fire an action potential. Together, synaptic summation and the threshold for excitation act as a filter so that random “noise” in the system is not transmitted as important information.
Everyday Connection: Brain-computer interface
Amyotrophic lateral sclerosis (ALS, also called Lou Gehrig’s Disease) is a neurological disease characterized by the degeneration of the motor neurons that control voluntary movements. The disease begins with muscle weakening and lack of coordination and eventually destroys the neurons that control speech, breathing, and swallowing; in the end, the disease can lead to paralysis. At that point, patients require assistance from machines to be able to breathe and to communicate. Several special technologies have been developed to allow “locked-in” patients to communicate with the rest of the world. One technology, for example, allows patients to type out sentences by twitching their cheek. These sentences can then be read aloud by a computer.
A relatively new line of research for helping paralyzed patients, including those with ALS, to communicate and retain a degree of self-sufficiency is called brain-computer interface (BCI) technology and is illustrated in Figure \(9\). This technology sounds like something out of science fiction: it allows paralyzed patients to control a computer using only their thoughts. There are several forms of BCI. Some forms use EEG recordings from electrodes taped onto the skull. These recordings contain information from large populations of neurons that can be decoded by a computer. Other forms of BCI require the implantation of an array of electrodes smaller than a postage stamp in the arm and hand area of the motor cortex. This form of BCI, while more invasive, is very powerful as each electrode can record actual action potentials from one or more neurons. These signals are then sent to a computer, which has been trained to decode the signal and feed it to a tool—such as a cursor on a computer screen. This means that a patient with ALS can use e-mail, read the Internet, and communicate with others by thinking of moving his or her hand or arm (even though the paralyzed patient cannot make that bodily movement). Recent advances have allowed a paralyzed locked-in patient who suffered a stroke 15 years ago to control a robotic arm and even to feed herself coffee using BCI technology.
Despite the amazing advancements in BCI technology, it also has limitations. The technology can require many hours of training and long periods of intense concentration for the patient; it can also require brain surgery to implant the devices.
Link to Learning
Watch this video in which a paralyzed woman use a brain-controlled robotic arm to bring a drink to her mouth, among other images of brain-computer interface technology in action.
Synaptic Plasticity
Synapses are not static structures. They can be weakened or strengthened. They can be broken, and new synapses can be made. Synaptic plasticity allows for these changes, which are all needed for a functioning nervous system. In fact, synaptic plasticity is the basis of learning and memory. Two processes in particular, long-term potentiation (LTP) and long-term depression (LTD) are important forms of synaptic plasticity that occur in synapses in the hippocampus, a brain region that is involved in storing memories.
Long-term Potentiation (LTP)
Long-term potentiation (LTP) is a persistent strengthening of a synaptic connection. LTP is based on the Hebbian principle: cells that fire together wire together. There are various mechanisms, none fully understood, behind the synaptic strengthening seen with LTP. One known mechanism involves a type of postsynaptic glutamate receptor, called NMDA (N-Methyl-D-aspartate) receptors, shown in Figure \(10\). These receptors are normally blocked by magnesium ions; however, when the postsynaptic neuron is depolarized by multiple presynaptic inputs in quick succession (either from one neuron or multiple neurons), the magnesium ions are forced out allowing Ca ions to pass into the postsynaptic cell. Next, Ca2+ ions entering the cell initiate a signaling cascade that causes a different type of glutamate receptor, called AMPA (α-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid) receptors, to be inserted into the postsynaptic membrane, since activated AMPA receptors allow positive ions to enter the cell. So, the next time glutamate is released from the presynaptic membrane, it will have a larger excitatory effect (EPSP) on the postsynaptic cell because the binding of glutamate to these AMPA receptors will allow more positive ions into the cell. The insertion of additional AMPA receptors strengthens the synapse and means that the postsynaptic neuron is more likely to fire in response to presynaptic neurotransmitter release. Some drugs of abuse co-opt the LTP pathway, and this synaptic strengthening can lead to addiction.
Long-term Depression (LTD)
Long-term depression (LTD) is essentially the reverse of LTP: it is a long-term weakening of a synaptic connection. One mechanism known to cause LTD also involves AMPA receptors. In this situation, calcium that enters through NMDA receptors initiates a different signaling cascade, which results in the removal of AMPA receptors from the postsynaptic membrane, as illustrated in Figure \(10\). The decrease in AMPA receptors in the membrane makes the postsynaptic neuron less responsive to glutamate released from the presynaptic neuron. While it may seem counterintuitive, LTD may be just as important for learning and memory as LTP. The weakening and pruning of unused synapses allows for unimportant connections to be lost and makes the synapses that have undergone LTP that much stronger by comparison.
Summary
Neurons have charged membranes because there are different concentrations of ions inside and outside of the cell. Voltage-gated ion channels control the movement of ions into and out of a neuron. When a neuronal membrane is depolarized to at least the threshold of excitation, an action potential is fired. The action potential is then propagated along a myelinated axon to the axon terminals. In a chemical synapse, the action potential causes release of neurotransmitter molecules into the synaptic cleft. Through binding to postsynaptic receptors, the neurotransmitter can cause excitatory or inhibitory postsynaptic potentials by depolarizing or hyperpolarizing, respectively, the postsynaptic membrane. In electrical synapses, the action potential is directly communicated to the postsynaptic cell through gap junctions—large channel proteins that connect the pre-and postsynaptic membranes. Synapses are not static structures and can be strengthened and weakened. Two mechanisms of synaptic plasticity are long-term potentiation and long-term depression.
Art Connections
Figure \(3\): Potassium channel blockers, such as amiodarone and procainamide, which are used to treat abnormal electrical activity in the heart, called cardiac dysrhythmia, impede the movement of K+ through voltage-gated K+ channels. Which part of the action potential would you expect potassium channels to affect?
Answer
Potassium channel blockers slow the repolarization phase, but have no effect on depolarization.
Glossary
action potential
self-propagating momentary change in the electrical potential of a neuron (or muscle) membrane
depolarization
change in the membrane potential to a less negative value
excitatory postsynaptic potential (EPSP)
depolarization of a postsynaptic membrane caused by neurotransmitter molecules released from a presynaptic cell
hyperpolarization
change in the membrane potential to a more negative value
inhibitory postsynaptic potential (IPSP)
hyperpolarization of a postsynaptic membrane caused by neurotransmitter molecules released from a presynaptic cell
long-term depression (LTD)
prolonged decrease in synaptic coupling between a pre- and postsynaptic cell
long-term potentiation (LTP)
prolonged increase in synaptic coupling between a pre-and postsynaptic cell
membrane potential
difference in electrical potential between the inside and outside of a cell
refractory period
period after an action potential when it is more difficult or impossible for an action potential to be fired; caused by inactivation of sodium channels and activation of additional potassium channels of the membrane
saltatory conduction
“jumping” of an action potential along an axon from one node of Ranvier to the next
summation
process of multiple presynaptic inputs creating EPSPs around the same time for the postsynaptic neuron to be sufficiently depolarized to fire an action potential
synaptic cleft
space between the presynaptic and postsynaptic membranes
synaptic vesicle
spherical structure that contains a neurotransmitter
threshold of excitation
level of depolarization needed for an action potential to fire
42.02: The Mechanism of Nerve Impulse Transmission
Skills to Develop
• Identify the general and special senses in humans
• Describe three important steps in sensory perception
• Explain the concept of just-noticeable difference in sensory perception
Senses provide information about the body and its environment. Humans have five special senses: olfaction (smell), gustation (taste), equilibrium (balance and body position), vision, and hearing. Additionally, we possess general senses, also called somatosensation, which respond to stimuli like temperature, pain, pressure, and vibration. Vestibular sensation, which is an organism’s sense of spatial orientation and balance, proprioception (position of bones, joints, and muscles), and the sense of limb position that is used to track kinesthesia (limb movement) are part of somatosensation. Although the sensory systems associated with these senses are very different, all share a common function: to convert a stimulus (such as light, or sound, or the position of the body) into an electrical signal in the nervous system. This process is called sensory transduction.
There are two broad types of cellular systems that perform sensory transduction. In one, a neuron works with a sensory receptor, a cell, or cell process that is specialized to engage with and detect a specific stimulus. Stimulation of the sensory receptor activates the associated afferent neuron, which carries information about the stimulus to the central nervous system. In the second type of sensory transduction, a sensory nerve ending responds to a stimulus in the internal or external environment: this neuron constitutes the sensory receptor. Free nerve endings can be stimulated by several different stimuli, thus showing little receptor specificity. For example, pain receptors in your gums and teeth may be stimulated by temperature changes, chemical stimulation, or pressure.
Reception
The first step in sensation is reception, which is the activation of sensory receptors by stimuli such as mechanical stimuli (being bent or squished, for example), chemicals, or temperature. The receptor can then respond to the stimuli. The region in space in which a given sensory receptor can respond to a stimulus, be it far away or in contact with the body, is that receptor’s receptive field. Think for a moment about the differences in receptive fields for the different senses. For the sense of touch, a stimulus must come into contact with body. For the sense of hearing, a stimulus can be a moderate distance away (some baleen whale sounds can propagate for many kilometers). For vision, a stimulus can be very far away; for example, the visual system perceives light from stars at enormous distances.
Transduction
The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system. This takes place at the sensory receptor, and the change in electrical potential that is produced is called the receptor potential. How is sensory input, such as pressure on the skin, changed to a receptor potential? In this example, a type of receptor called a mechanoreceptor (as shown in Figure \(1\)) possesses specialized membranes that respond to pressure. Disturbance of these dendrites by compressing them or bending them opens gated ion channels in the plasma membrane of the sensory neuron, changing its electrical potential. Recall that in the nervous system, a positive change of a neuron’s electrical potential (also called the membrane potential), depolarizes the neuron. Receptor potentials are graded potentials: the magnitude of these graded (receptor) potentials varies with the strength of the stimulus. If the magnitude of depolarization is sufficient (that is, if membrane potential reaches a threshold), the neuron will fire an action potential. In most cases, the correct stimulus impinging on a sensory receptor will drive membrane potential in a positive direction, although for some receptors, such as those in the visual system, this is not always the case.
Sensory receptors for different senses are very different from each other, and they are specialized according to the type of stimulus they sense: they have receptor specificity. For example, touch receptors, light receptors, and sound receptors are each activated by different stimuli. Touch receptors are not sensitive to light or sound; they are sensitive only to touch or pressure. However, stimuli may be combined at higher levels in the brain, as happens with olfaction, contributing to our sense of taste.
Encoding and Transmission of Sensory Information
Four aspects of sensory information are encoded by sensory systems: the type of stimulus, the location of the stimulus in the receptive field, the duration of the stimulus, and the relative intensity of the stimulus. Thus, action potentials transmitted over a sensory receptor’s afferent axons encode one type of stimulus, and this segregation of the senses is preserved in other sensory circuits. For example, auditory receptors transmit signals over their own dedicated system, and electrical activity in the axons of the auditory receptors will be interpreted by the brain as an auditory stimulus—a sound.
The intensity of a stimulus is often encoded in the rate of action potentials produced by the sensory receptor. Thus, an intense stimulus will produce a more rapid train of action potentials, and reducing the stimulus will likewise slow the rate of production of action potentials. A second way in which intensity is encoded is by the number of receptors activated. An intense stimulus might initiate action potentials in a large number of adjacent receptors, while a less intense stimulus might stimulate fewer receptors. Integration of sensory information begins as soon as the information is received in the CNS, and the brain will further process incoming signals.
Perception
Perception is an individual’s interpretation of a sensation. Although perception relies on the activation of sensory receptors, perception happens not at the level of the sensory receptor, but at higher levels in the nervous system, in the brain. The brain distinguishes sensory stimuli through a sensory pathway: action potentials from sensory receptors travel along neurons that are dedicated to a particular stimulus. These neurons are dedicated to that particular stimulus and synapse with particular neurons in the brain or spinal cord.
All sensory signals, except those from the olfactory system, are transmitted though the central nervous system and are routed to the thalamus and to the appropriate region of the cortex. Recall that the thalamus is a structure in the forebrain that serves as a clearinghouse and relay station for sensory (as well as motor) signals. When the sensory signal exits the thalamus, it is conducted to the specific area of the cortex (Figure \(2\)) dedicated to processing that particular sense.
How are neural signals interpreted? Interpretation of sensory signals between individuals of the same species is largely similar, owing to the inherited similarity of their nervous systems; however, there are some individual differences. A good example of this is individual tolerances to a painful stimulus, such as dental pain, which certainly differ.
Scientific Method Connection: Just-Noticeable Difference
It is easy to differentiate between a one-pound bag of rice and a two-pound bag of rice. There is a one-pound difference, and one bag is twice as heavy as the other. However, would it be as easy to differentiate between a 20- and a 21-pound bag?
Question: What is the smallest detectible weight difference between a one-pound bag of rice and a larger bag? What is the smallest detectible difference between a 20-pound bag and a larger bag? In both cases, at what weights are the differences detected? This smallest detectible difference in stimuli is known as the just-noticeable difference (JND).
Background: Research background literature on JND and on Weber’s Law, a description of a proposed mathematical relationship between the overall magnitude of the stimulus and the JND. You will be testing JND of different weights of rice in bags. Choose a convenient increment that is to be stepped through while testing. For example, you could choose 10 percent increments between one and two pounds (1.1, 1.2, 1.3, 1.4, and so on) or 20 percent increments (1.2, 1.4, 1.6, and 1.8).
Hypothesis: Develop a hypothesis about JND in terms of percentage of the whole weight being tested (such as “the JND between the two small bags and between the two large bags is proportionally the same,” or “. . . is not proportionally the same.”) So, for the first hypothesis, if the JND between the one-pound bag and a larger bag is 0.2 pounds (that is, 20 percent; 1.0 pound feels the same as 1.1 pounds, but 1.0 pound feels less than 1.2 pounds), then the JND between the 20-pound bag and a larger bag will also be 20 percent. (So, 20 pounds feels the same as 22 pounds or 23 pounds, but 20 pounds feels less than 24 pounds.)
Test the hypothesis: Enlist 24 participants, and split them into two groups of 12. To set up the demonstration, assuming a 10 percent increment was selected, have the first group be the one-pound group. As a counter-balancing measure against a systematic error, however, six of the first group will compare one pound to two pounds, and step down in weight (1.0 to 2.0, 1.0 to 1.9, and so on.), while the other six will step up (1.0 to 1.1, 1.0 to 1.2, and so on). Apply the same principle to the 20-pound group (20 to 40, 20 to 38, and so on, and 20 to 22, 20 to 24, and so on). Given the large difference between 20 and 40 pounds, you may wish to use 30 pounds as your larger weight. In any case, use two weights that are easily detectable as different.
Record the observations: Record the data in a table similar to the table below. For the one-pound and 20-pound groups (base weights) record a plus sign (+) for each participant that detects a difference between the base weight and the step weight. Record a minus sign (-) for each participant that finds no difference. If one-tenth steps were not used, then replace the steps in the “Step Weight” columns with the step you are using.
Table \(1\): Results of JND Testing (+ = difference; – = no difference)
Step Weight One pound 20 pounds Step Weight
1.1 22
1.2 24
1.3 26
1.4 28
1.5 30
1.6 32
1.7 34
1.8 36
1.9 38
2.0 40
Analyze the data/report the results: What step weight did all participants find to be equal with one-pound base weight? What about the 20-pound group?
Draw a conclusion: Did the data support the hypothesis? Are the final weights proportionally the same? If not, why not? Do the findings adhere to Weber’s Law? Weber’s Law states that the concept that a just-noticeable difference in a stimulus is proportional to the magnitude of the original stimulus.
Summary
A sensory activation occurs when a physical or chemical stimulus is processed into a neural signal (sensory transduction) by a sensory receptor. Perception is an individual interpretation of a sensation and is a brain function. Humans have special senses: olfaction, gustation, equilibrium, and hearing, plus the general senses of somatosensation.
Sensory receptors are either specialized cells associated with sensory neurons or the specialized ends of sensory neurons that are a part of the peripheral nervous system, and they are used to receive information about the environment (internal or external). Each sensory receptor is modified for the type of stimulus it detects. For example, neither gustatory receptors nor auditory receptors are sensitive to light. Each sensory receptor is responsive to stimuli within a specific region in space, which is known as that receptor’s receptive field. The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system.
All sensory signals, except those from the olfactory system, enter the central nervous system and are routed to the thalamus. When the sensory signal exits the thalamus, it is conducted to the specific area of the cortex dedicated to processing that particular sense.
Glossary
kinesthesia
sense of body movement
mechanoreceptor
sensory receptor modified to respond to mechanical disturbance such as being bent, touch, pressure, motion, and sound
perception
individual interpretation of a sensation; a brain function
proprioception
sense of limb position; used to track kinesthesia
reception
receipt of a signal (such as light or sound) by sensory receptors
receptive field
region in space in which a stimulus can activate a given sensory receptor
receptor potential
membrane potential in a sensory receptor in response to detection of a stimulus
sensory receptor
specialized neuron or other cells associated with a neuron that is modified to receive specific sensory input
sensory transduction
conversion of a sensory stimulus into electrical energy in the nervous system by a change in the membrane potential
vestibular sense
sense of spatial orientation and balance | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/42%3A_The_Nervous_System/42.02%3A_The_Mechanism_of_Nerve_Impulse_Transmission/42.2.01%3A_Sensory_Processes.txt |
Skills to Develop
• Identify the spinal cord, cerebral lobes, and other brain areas on a diagram of the brain
• Describe the basic functions of the spinal cord, cerebral lobes, and other brain areas
The central nervous system (CNS) is made up of the brain, a part of which is shown in Figure \(1\) and spinal cord and is covered with three layers of protective coverings called meninges (from the Greek word for membrane). The outermost layer is the dura mater (Latin for “hard mother”). As the Latin suggests, the primary function for this thick layer is to protect the brain and spinal cord. The dura mater also contains vein-like structures that carry blood from the brain back to the heart. The middle layer is the web-like arachnoid mater. The last layer is the pia mater (Latin for “soft mother”), which directly contacts and covers the brain and spinal cord like plastic wrap. The space between the arachnoid and pia maters is filled with cerebrospinal fluid (CSF). CSF is produced by a tissue called choroid plexus in fluid-filled compartments in the CNS called ventricles. The brain floats in CSF, which acts as a cushion and shock absorber and makes the brain neutrally buoyant. CSF also functions to circulate chemical substances throughout the brain and into the spinal cord.
The entire brain contains only about 8.5 tablespoons of CSF, but CSF is constantly produced in the ventricles. This creates a problem when a ventricle is blocked—the CSF builds up and creates swelling and the brain is pushed against the skull. This swelling condition is called hydrocephalus (“water head”) and can cause seizures, cognitive problems, and even death if a shunt is not inserted to remove the fluid and pressure.
Brain
The brain is the part of the central nervous system that is contained in the cranial cavity of the skull. It includes the cerebral cortex, limbic system, basal ganglia, thalamus, hypothalamus, and cerebellum. There are three different ways that a brain can be sectioned in order to view internal structures: a sagittal section cuts the brain left to right, as shown in Figure \(2\)b, a coronal section cuts the brain front to back, as shown in Figure \(2\)a, and a horizontal section cuts the brain top to bottom.
Cerebral Cortex
The outermost part of the brain is a thick piece of nervous system tissue called the cerebral cortex, which is folded into hills called gyri (singular: gyrus) and valleys called sulci (singular: sulcus). The cortex is made up of two hemispheres—right and left—which are separated by a large sulcus. A thick fiber bundle called the corpus callosum (Latin: “tough body”) connects the two hemispheres and allows information to be passed from one side to the other. Although there are some brain functions that are localized more to one hemisphere than the other, the functions of the two hemispheres are largely redundant. In fact, sometimes (very rarely) an entire hemisphere is removed to treat severe epilepsy. While patients do suffer some deficits following the surgery, they can have surprisingly few problems, especially when the surgery is performed on children who have very immature nervous systems.
In other surgeries to treat severe epilepsy, the corpus callosum is cut instead of removing an entire hemisphere. This causes a condition called split-brain, which gives insights into unique functions of the two hemispheres. For example, when an object is presented to patients’ left visual field, they may be unable to verbally name the object (and may claim to not have seen an object at all). This is because the visual input from the left visual field crosses and enters the right hemisphere and cannot then signal to the speech center, which generally is found in the left side of the brain. Remarkably, if a split-brain patient is asked to pick up a specific object out of a group of objects with the left hand, the patient will be able to do so but will still be unable to vocally identify it.
Link to Learning
See this website to learn more about split-brain patients and to play a game where you can model the split-brain experiments yourself.
Each cortical hemisphere contains regions called lobes that are involved in different functions. Scientists use various techniques to determine what brain areas are involved in different functions: they examine patients who have had injuries or diseases that affect specific areas and see how those areas are related to functional deficits. They also conduct animal studies where they stimulate brain areas and see if there are any behavioral changes. They use a technique called transmagnetic stimulation (TMS) to temporarily deactivate specific parts of the cortex using strong magnets placed outside the head; and they use functional magnetic resonance imaging (fMRI) to look at changes in oxygenated blood flow in particular brain regions that correlate with specific behavioral tasks. These techniques, and others, have given great insight into the functions of different brain regions but have also showed that any given brain area can be involved in more than one behavior or process, and any given behavior or process generally involves neurons in multiple brain areas. That being said, each hemisphere of the mammalian cerebral cortex can be broken down into four functionally and spatially defined lobes: frontal, parietal, temporal, and occipital. Figure \(3\) illustrates these four lobes of the human cerebral cortex.
The frontal lobe is located at the front of the brain, over the eyes. This lobe contains the olfactory bulb, which processes smells. The frontal lobe also contains the motor cortex, which is important for planning and implementing movement. Areas within the motor cortex map to different muscle groups, and there is some organization to this map, as shown in Figure \(4\). For example, the neurons that control movement of the fingers are next to the neurons that control movement of the hand. Neurons in the frontal lobe also control cognitive functions like maintaining attention, speech, and decision-making. Studies of humans who have damaged their frontal lobes show that parts of this area are involved in personality, socialization, and assessing risk.
The parietal lobe is located at the top of the brain. Neurons in the parietal lobe are involved in speech and also reading. Two of the parietal lobe’s main functions are processing somatosensation—touch sensations like pressure, pain, heat, cold—and processing proprioception—the sense of how parts of the body are oriented in space. The parietal lobe contains a somatosensory map of the body similar to the motor cortex.
The occipital lobe is located at the back of the brain. It is primarily involved in vision—seeing, recognizing, and identifying the visual world.
The temporal lobe is located at the base of the brain by your ears and is primarily involved in processing and interpreting sounds. It also contains the hippocampus (Greek for “seahorse”)—a structure that processes memory formation. The hippocampus is illustrated in Figure \(6\). The role of the hippocampus in memory was partially determined by studying one famous epileptic patient, HM, who had both sides of his hippocampus removed in an attempt to cure his epilepsy. His seizures went away, but he could no longer form new memories (although he could remember some facts from before his surgery and could learn new motor tasks).
Evolution Connection: Cerebral Cortex
Compared to other vertebrates, mammals have exceptionally large brains for their body size. An entire alligator’s brain, for example, would fill about one and a half teaspoons. This increase in brain to body size ratio is especially pronounced in apes, whales, and dolphins. While this increase in overall brain size doubtlessly played a role in the evolution of complex behaviors unique to mammals, it does not tell the whole story. Scientists have found a relationship between the relatively high surface area of the cortex and the intelligence and complex social behaviors exhibited by some mammals. This increased surface area is due, in part, to increased folding of the cortical sheet (more sulci and gyri). For example, a rat cortex is very smooth with very few sulci and gyri. Cat and sheep cortices have more sulci and gyri. Chimps, humans, and dolphins have even more.
Basal Ganglia
Interconnected brain areas called the basal ganglia (or basal nuclei) play important roles in movement control and posture. Damage to the basal ganglia, as in Parkinson’s disease, leads to motor impairments like a shuffling gait when walking. The basal ganglia also regulate motivation. For example, when a wasp sting led to bilateral basal ganglia damage in a 25-year-old businessman, he began to spend all his days in bed and showed no interest in anything or anybody. But when he was externally stimulated—as when someone asked to play a card game with him—he was able to function normally. Interestingly, he and other similar patients do not report feeling bored or frustrated by their state.
Thalamus
The thalamus (Greek for “inner chamber”), illustrated in Figure \(6\), acts as a gateway to and from the cortex. It receives sensory and motor inputs from the body and also receives feedback from the cortex. This feedback mechanism can modulate conscious awareness of sensory and motor inputs depending on the attention and arousal state of the animal. The thalamus helps regulate consciousness, arousal, and sleep states. A rare genetic disorder called fatal familial insomnia causes the degeneration of thalamic neurons and glia. This disorder prevents affected patients from being able to sleep, among other symptoms, and is eventually fatal.
Hypothalamus
Below the thalamus is the hypothalamus, shown in Figure \(6\). The hypothalamus controls the endocrine system by sending signals to the pituitary gland, a pea-sized endocrine gland that releases several different hormones that affect other glands as well as other cells. This relationship means that the hypothalamus regulates important behaviors that are controlled by these hormones. The hypothalamus is the body’s thermostat—it makes sure key functions like food and water intake, energy expenditure, and body temperature are kept at appropriate levels. Neurons within the hypothalamus also regulate circadian rhythms, sometimes called sleep cycles.
Limbic System
The limbic system is a connected set of structures that regulates emotion, as well as behaviors related to fear and motivation. It plays a role in memory formation and includes parts of the thalamus and hypothalamus as well as the hippocampus. One important structure within the limbic system is a temporal lobe structure called the amygdala (Greek for “almond”), illustrated in Figure \(6\). The two amygdala are important both for the sensation of fear and for recognizing fearful faces. The cingulate gyrus helps regulate emotions and pain.
Cerebellum
The cerebellum (Latin for “little brain”), shown in Figure \(3\), sits at the base of the brain on top of the brainstem. The cerebellum controls balance and aids in coordinating movement and learning new motor tasks.
Brainstem
The brainstem, illustrated in Figure \(3\), connects the rest of the brain with the spinal cord. It consists of the midbrain, medulla oblongata, and the pons. Motor and sensory neurons extend through the brainstem allowing for the relay of signals between the brain and spinal cord. Ascending neural pathways cross in this section of the brain allowing the left hemisphere of the cerebrum to control the right side of the body and vice versa. The brainstem coordinates motor control signals sent from the brain to the body. The brainstem controls several important functions of the body including alertness, arousal, breathing, blood pressure, digestion, heart rate, swallowing, walking, and sensory and motor information integration.
Spinal Cord
Connecting to the brainstem and extending down the body through the spinal column is the spinal cord, shown in Figure \(3\). The spinal cord is a thick bundle of nerve tissue that carries information about the body to the brain and from the brain to the body. The spinal cord is contained within the bones of the vertebrate column but is able to communicate signals to and from the body through its connections with spinal nerves (part of the peripheral nervous system). A cross-section of the spinal cord looks like a white oval containing a gray butterfly-shape, as illustrated in Figure \(7\). Myelinated axons make up the “white matter” and neuron and glial cell bodies make up the “gray matter.” Gray matter is also composed of interneurons, which connect two neurons each located in different parts of the body. Axons and cell bodies in the dorsal (facing the back of the animal) spinal cord convey mostly sensory information from the body to the brain. Axons and cell bodies in the ventral (facing the front of the animal) spinal cord primarily transmit signals controlling movement from the brain to the body.
The spinal cord also controls motor reflexes. These reflexes are quick, unconscious movements—like automatically removing a hand from a hot object. Reflexes are so fast because they involve local synaptic connections. For example, the knee reflex that a doctor tests during a routine physical is controlled by a single synapse between a sensory neuron and a motor neuron. While a reflex may only require the involvement of one or two synapses, synapses with interneurons in the spinal column transmit information to the brain to convey what happened (the knee jerked, or the hand was hot).
In the United States, there around 10,000 spinal cord injuries each year. Because the spinal cord is the information superhighway connecting the brain with the body, damage to the spinal cord can lead to paralysis. The extent of the paralysis depends on the location of the injury along the spinal cord and whether the spinal cord was completely severed. For example, if the spinal cord is damaged at the level of the neck, it can cause paralysis from the neck down, whereas damage to the spinal column further down may limit paralysis to the legs. Spinal cord injuries are notoriously difficult to treat because spinal nerves do not regenerate, although ongoing research suggests that stem cell transplants may be able to act as a bridge to reconnect severed nerves. Researchers are also looking at ways to prevent the inflammation that worsens nerve damage after injury. One such treatment is to pump the body with cold saline to induce hypothermia. This cooling can prevent swelling and other processes that are thought to worsen spinal cord injuries.
Summary
The vertebrate central nervous system contains the brain and the spinal cord, which are covered and protected by three meninges. The brain contains structurally and functionally defined regions. In mammals, these include the cortex (which can be broken down into four primary functional lobes: frontal, temporal, occipital, and parietal), basal ganglia, thalamus, hypothalamus, limbic system, cerebellum, and brainstem—although structures in some of these designations overlap. While functions may be primarily localized to one structure in the brain, most complex functions, like language and sleep, involve neurons in multiple brain regions. The spinal cord is the information superhighway that connects the brain with the rest of the body through its connections with peripheral nerves. It transmits sensory and motor input and also controls motor reflexes.
Glossary
amygdala
structure within the limbic system that processes fear
arachnoid mater
spiderweb-like middle layer of the meninges that cover the central nervous system
basal ganglia
interconnected collections of cells in the brain that are involved in movement and motivation; also known as basal nuclei
basal nuclei
see basal ganglia
brainstem
portion of the brain that connects with the spinal cord; controls basic nervous system functions like breathing, heart rate, and swallowing
cerebellum
brain structure involved in posture, motor coordination, and learning new motor actions
cerebral cortex
outermost sheet of brain tissue; involved in many higher-order functions
choroid plexus
spongy tissue within ventricles that produces cerebrospinal fluid
cingulate gyrus
helps regulate emotions and pain; thought to directly drive the body’s conscious response to unpleasant experiences
corpus callosum
thick fiber bundle that connects the cerebral hemispheres
cerebrospinal fluid (CSF)
clear liquid that surrounds the brain and spinal cord and fills the ventricles and central canal; acts as a shock absorber and circulates material throughout the brain and spinal cord.
dura mater
tough outermost layer that covers the central nervous system
frontal lobe
part of the cerebral cortex that contains the motor cortex and areas involved in planning, attention, and language
gyrus
(plural: gyri) ridged protrusions in the cortex
hippocampus
brain structure in the temporal lobe involved in processing memories
hypothalamus
brain structure that controls hormone release and body homeostasis
limbic system
connected brain areas that process emotion and motivation
meninge
membrane that covers and protects the central nervous system
occipital lobe
part of the cerebral cortex that contains visual cortex and processes visual stimuli
parietal lobe
part of the cerebral cortex involved in processing touch and the sense of the body in space
pia mater
thin membrane layer directly covering the brain and spinal cord
proprioception
sense about how parts of the body are oriented in space
somatosensation
sense of touch
spinal cord
thick fiber bundle that connects the brain with peripheral nerves; transmits sensory and motor information; contains neurons that control motor reflexes
sulcus
(plural: sulci) indents or “valleys” in the cortex
temporal lobe
part of the cerebral cortex that processes auditory input; parts of the temporal lobe are involved in speech, memory, and emotion processing
thalamus
brain area that relays sensory information to the cortex
ventricle
cavity within brain that contains cerebrospinal fluid | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/42%3A_The_Nervous_System/42.03%3A_Synapses-_Where_Neurons_Communicate_With_Other_Cells.txt |
Skills to Develop
• Identify the spinal cord, cerebral lobes, and other brain areas on a diagram of the brain
• Describe the basic functions of the spinal cord, cerebral lobes, and other brain areas
The central nervous system (CNS) is made up of the brain, a part of which is shown in Figure \(1\) and spinal cord and is covered with three layers of protective coverings called meninges (from the Greek word for membrane). The outermost layer is the dura mater (Latin for “hard mother”). As the Latin suggests, the primary function for this thick layer is to protect the brain and spinal cord. The dura mater also contains vein-like structures that carry blood from the brain back to the heart. The middle layer is the web-like arachnoid mater. The last layer is the pia mater (Latin for “soft mother”), which directly contacts and covers the brain and spinal cord like plastic wrap. The space between the arachnoid and pia maters is filled with cerebrospinal fluid (CSF). CSF is produced by a tissue called choroid plexus in fluid-filled compartments in the CNS called ventricles. The brain floats in CSF, which acts as a cushion and shock absorber and makes the brain neutrally buoyant. CSF also functions to circulate chemical substances throughout the brain and into the spinal cord.
The entire brain contains only about 8.5 tablespoons of CSF, but CSF is constantly produced in the ventricles. This creates a problem when a ventricle is blocked—the CSF builds up and creates swelling and the brain is pushed against the skull. This swelling condition is called hydrocephalus (“water head”) and can cause seizures, cognitive problems, and even death if a shunt is not inserted to remove the fluid and pressure.
Brain
The brain is the part of the central nervous system that is contained in the cranial cavity of the skull. It includes the cerebral cortex, limbic system, basal ganglia, thalamus, hypothalamus, and cerebellum. There are three different ways that a brain can be sectioned in order to view internal structures: a sagittal section cuts the brain left to right, as shown in Figure \(2\)b, a coronal section cuts the brain front to back, as shown in Figure \(2\)a, and a horizontal section cuts the brain top to bottom.
Cerebral Cortex
The outermost part of the brain is a thick piece of nervous system tissue called the cerebral cortex, which is folded into hills called gyri (singular: gyrus) and valleys called sulci (singular: sulcus). The cortex is made up of two hemispheres—right and left—which are separated by a large sulcus. A thick fiber bundle called the corpus callosum (Latin: “tough body”) connects the two hemispheres and allows information to be passed from one side to the other. Although there are some brain functions that are localized more to one hemisphere than the other, the functions of the two hemispheres are largely redundant. In fact, sometimes (very rarely) an entire hemisphere is removed to treat severe epilepsy. While patients do suffer some deficits following the surgery, they can have surprisingly few problems, especially when the surgery is performed on children who have very immature nervous systems.
In other surgeries to treat severe epilepsy, the corpus callosum is cut instead of removing an entire hemisphere. This causes a condition called split-brain, which gives insights into unique functions of the two hemispheres. For example, when an object is presented to patients’ left visual field, they may be unable to verbally name the object (and may claim to not have seen an object at all). This is because the visual input from the left visual field crosses and enters the right hemisphere and cannot then signal to the speech center, which generally is found in the left side of the brain. Remarkably, if a split-brain patient is asked to pick up a specific object out of a group of objects with the left hand, the patient will be able to do so but will still be unable to vocally identify it.
Link to Learning
See this website to learn more about split-brain patients and to play a game where you can model the split-brain experiments yourself.
Each cortical hemisphere contains regions called lobes that are involved in different functions. Scientists use various techniques to determine what brain areas are involved in different functions: they examine patients who have had injuries or diseases that affect specific areas and see how those areas are related to functional deficits. They also conduct animal studies where they stimulate brain areas and see if there are any behavioral changes. They use a technique called transmagnetic stimulation (TMS) to temporarily deactivate specific parts of the cortex using strong magnets placed outside the head; and they use functional magnetic resonance imaging (fMRI) to look at changes in oxygenated blood flow in particular brain regions that correlate with specific behavioral tasks. These techniques, and others, have given great insight into the functions of different brain regions but have also showed that any given brain area can be involved in more than one behavior or process, and any given behavior or process generally involves neurons in multiple brain areas. That being said, each hemisphere of the mammalian cerebral cortex can be broken down into four functionally and spatially defined lobes: frontal, parietal, temporal, and occipital. Figure \(3\) illustrates these four lobes of the human cerebral cortex.
The frontal lobe is located at the front of the brain, over the eyes. This lobe contains the olfactory bulb, which processes smells. The frontal lobe also contains the motor cortex, which is important for planning and implementing movement. Areas within the motor cortex map to different muscle groups, and there is some organization to this map, as shown in Figure \(4\). For example, the neurons that control movement of the fingers are next to the neurons that control movement of the hand. Neurons in the frontal lobe also control cognitive functions like maintaining attention, speech, and decision-making. Studies of humans who have damaged their frontal lobes show that parts of this area are involved in personality, socialization, and assessing risk.
The parietal lobe is located at the top of the brain. Neurons in the parietal lobe are involved in speech and also reading. Two of the parietal lobe’s main functions are processing somatosensation—touch sensations like pressure, pain, heat, cold—and processing proprioception—the sense of how parts of the body are oriented in space. The parietal lobe contains a somatosensory map of the body similar to the motor cortex.
The occipital lobe is located at the back of the brain. It is primarily involved in vision—seeing, recognizing, and identifying the visual world.
The temporal lobe is located at the base of the brain by your ears and is primarily involved in processing and interpreting sounds. It also contains the hippocampus (Greek for “seahorse”)—a structure that processes memory formation. The hippocampus is illustrated in Figure \(6\). The role of the hippocampus in memory was partially determined by studying one famous epileptic patient, HM, who had both sides of his hippocampus removed in an attempt to cure his epilepsy. His seizures went away, but he could no longer form new memories (although he could remember some facts from before his surgery and could learn new motor tasks).
Evolution Connection: Cerebral Cortex
Compared to other vertebrates, mammals have exceptionally large brains for their body size. An entire alligator’s brain, for example, would fill about one and a half teaspoons. This increase in brain to body size ratio is especially pronounced in apes, whales, and dolphins. While this increase in overall brain size doubtlessly played a role in the evolution of complex behaviors unique to mammals, it does not tell the whole story. Scientists have found a relationship between the relatively high surface area of the cortex and the intelligence and complex social behaviors exhibited by some mammals. This increased surface area is due, in part, to increased folding of the cortical sheet (more sulci and gyri). For example, a rat cortex is very smooth with very few sulci and gyri. Cat and sheep cortices have more sulci and gyri. Chimps, humans, and dolphins have even more.
Basal Ganglia
Interconnected brain areas called the basal ganglia (or basal nuclei) play important roles in movement control and posture. Damage to the basal ganglia, as in Parkinson’s disease, leads to motor impairments like a shuffling gait when walking. The basal ganglia also regulate motivation. For example, when a wasp sting led to bilateral basal ganglia damage in a 25-year-old businessman, he began to spend all his days in bed and showed no interest in anything or anybody. But when he was externally stimulated—as when someone asked to play a card game with him—he was able to function normally. Interestingly, he and other similar patients do not report feeling bored or frustrated by their state.
Thalamus
The thalamus (Greek for “inner chamber”), illustrated in Figure \(6\), acts as a gateway to and from the cortex. It receives sensory and motor inputs from the body and also receives feedback from the cortex. This feedback mechanism can modulate conscious awareness of sensory and motor inputs depending on the attention and arousal state of the animal. The thalamus helps regulate consciousness, arousal, and sleep states. A rare genetic disorder called fatal familial insomnia causes the degeneration of thalamic neurons and glia. This disorder prevents affected patients from being able to sleep, among other symptoms, and is eventually fatal.
Hypothalamus
Below the thalamus is the hypothalamus, shown in Figure \(6\). The hypothalamus controls the endocrine system by sending signals to the pituitary gland, a pea-sized endocrine gland that releases several different hormones that affect other glands as well as other cells. This relationship means that the hypothalamus regulates important behaviors that are controlled by these hormones. The hypothalamus is the body’s thermostat—it makes sure key functions like food and water intake, energy expenditure, and body temperature are kept at appropriate levels. Neurons within the hypothalamus also regulate circadian rhythms, sometimes called sleep cycles.
Limbic System
The limbic system is a connected set of structures that regulates emotion, as well as behaviors related to fear and motivation. It plays a role in memory formation and includes parts of the thalamus and hypothalamus as well as the hippocampus. One important structure within the limbic system is a temporal lobe structure called the amygdala (Greek for “almond”), illustrated in Figure \(6\). The two amygdala are important both for the sensation of fear and for recognizing fearful faces. The cingulate gyrus helps regulate emotions and pain.
Cerebellum
The cerebellum (Latin for “little brain”), shown in Figure \(3\), sits at the base of the brain on top of the brainstem. The cerebellum controls balance and aids in coordinating movement and learning new motor tasks.
Brainstem
The brainstem, illustrated in Figure \(3\), connects the rest of the brain with the spinal cord. It consists of the midbrain, medulla oblongata, and the pons. Motor and sensory neurons extend through the brainstem allowing for the relay of signals between the brain and spinal cord. Ascending neural pathways cross in this section of the brain allowing the left hemisphere of the cerebrum to control the right side of the body and vice versa. The brainstem coordinates motor control signals sent from the brain to the body. The brainstem controls several important functions of the body including alertness, arousal, breathing, blood pressure, digestion, heart rate, swallowing, walking, and sensory and motor information integration.
Spinal Cord
Connecting to the brainstem and extending down the body through the spinal column is the spinal cord, shown in Figure \(3\). The spinal cord is a thick bundle of nerve tissue that carries information about the body to the brain and from the brain to the body. The spinal cord is contained within the bones of the vertebrate column but is able to communicate signals to and from the body through its connections with spinal nerves (part of the peripheral nervous system). A cross-section of the spinal cord looks like a white oval containing a gray butterfly-shape, as illustrated in Figure \(7\). Myelinated axons make up the “white matter” and neuron and glial cell bodies make up the “gray matter.” Gray matter is also composed of interneurons, which connect two neurons each located in different parts of the body. Axons and cell bodies in the dorsal (facing the back of the animal) spinal cord convey mostly sensory information from the body to the brain. Axons and cell bodies in the ventral (facing the front of the animal) spinal cord primarily transmit signals controlling movement from the brain to the body.
The spinal cord also controls motor reflexes. These reflexes are quick, unconscious movements—like automatically removing a hand from a hot object. Reflexes are so fast because they involve local synaptic connections. For example, the knee reflex that a doctor tests during a routine physical is controlled by a single synapse between a sensory neuron and a motor neuron. While a reflex may only require the involvement of one or two synapses, synapses with interneurons in the spinal column transmit information to the brain to convey what happened (the knee jerked, or the hand was hot).
In the United States, there around 10,000 spinal cord injuries each year. Because the spinal cord is the information superhighway connecting the brain with the body, damage to the spinal cord can lead to paralysis. The extent of the paralysis depends on the location of the injury along the spinal cord and whether the spinal cord was completely severed. For example, if the spinal cord is damaged at the level of the neck, it can cause paralysis from the neck down, whereas damage to the spinal column further down may limit paralysis to the legs. Spinal cord injuries are notoriously difficult to treat because spinal nerves do not regenerate, although ongoing research suggests that stem cell transplants may be able to act as a bridge to reconnect severed nerves. Researchers are also looking at ways to prevent the inflammation that worsens nerve damage after injury. One such treatment is to pump the body with cold saline to induce hypothermia. This cooling can prevent swelling and other processes that are thought to worsen spinal cord injuries.
Summary
The vertebrate central nervous system contains the brain and the spinal cord, which are covered and protected by three meninges. The brain contains structurally and functionally defined regions. In mammals, these include the cortex (which can be broken down into four primary functional lobes: frontal, temporal, occipital, and parietal), basal ganglia, thalamus, hypothalamus, limbic system, cerebellum, and brainstem—although structures in some of these designations overlap. While functions may be primarily localized to one structure in the brain, most complex functions, like language and sleep, involve neurons in multiple brain regions. The spinal cord is the information superhighway that connects the brain with the rest of the body through its connections with peripheral nerves. It transmits sensory and motor input and also controls motor reflexes.
Glossary
amygdala
structure within the limbic system that processes fear
arachnoid mater
spiderweb-like middle layer of the meninges that cover the central nervous system
basal ganglia
interconnected collections of cells in the brain that are involved in movement and motivation; also known as basal nuclei
basal nuclei
see basal ganglia
brainstem
portion of the brain that connects with the spinal cord; controls basic nervous system functions like breathing, heart rate, and swallowing
cerebellum
brain structure involved in posture, motor coordination, and learning new motor actions
cerebral cortex
outermost sheet of brain tissue; involved in many higher-order functions
choroid plexus
spongy tissue within ventricles that produces cerebrospinal fluid
cingulate gyrus
helps regulate emotions and pain; thought to directly drive the body’s conscious response to unpleasant experiences
corpus callosum
thick fiber bundle that connects the cerebral hemispheres
cerebrospinal fluid (CSF)
clear liquid that surrounds the brain and spinal cord and fills the ventricles and central canal; acts as a shock absorber and circulates material throughout the brain and spinal cord.
dura mater
tough outermost layer that covers the central nervous system
frontal lobe
part of the cerebral cortex that contains the motor cortex and areas involved in planning, attention, and language
gyrus
(plural: gyri) ridged protrusions in the cortex
hippocampus
brain structure in the temporal lobe involved in processing memories
hypothalamus
brain structure that controls hormone release and body homeostasis
limbic system
connected brain areas that process emotion and motivation
meninge
membrane that covers and protects the central nervous system
occipital lobe
part of the cerebral cortex that contains visual cortex and processes visual stimuli
parietal lobe
part of the cerebral cortex involved in processing touch and the sense of the body in space
pia mater
thin membrane layer directly covering the brain and spinal cord
proprioception
sense about how parts of the body are oriented in space
somatosensation
sense of touch
spinal cord
thick fiber bundle that connects the brain with peripheral nerves; transmits sensory and motor information; contains neurons that control motor reflexes
sulcus
(plural: sulci) indents or “valleys” in the cortex
temporal lobe
part of the cerebral cortex that processes auditory input; parts of the temporal lobe are involved in speech, memory, and emotion processing
thalamus
brain area that relays sensory information to the cortex
ventricle
cavity within brain that contains cerebrospinal fluid | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/42%3A_The_Nervous_System/42.04%3A_The_Central_Nervous_System-_Brain_and_Spinal_Cord.txt |
Skills to Develop
• Describe the organization and functions of the sympathetic and parasympathetic nervous systems
• Describe the organization and function of the sensory-somatic nervous system
The peripheral nervous system (PNS) is the connection between the central nervous system and the rest of the body. The CNS is like the power plant of the nervous system. It creates the signals that control the functions of the body. The PNS is like the wires that go to individual houses. Without those “wires,” the signals produced by the CNS could not control the body (and the CNS would not be able to receive sensory information from the body either).
The PNS can be broken down into the autonomic nervous system, which controls bodily functions without conscious control, and the sensory-somatic nervous system, which transmits sensory information from the skin, muscles, and sensory organs to the CNS and sends motor commands from the CNS to the muscles.
Autonomic Nervous System
Art Connection
Which of the following statements is false?
1. The parasympathetic pathway is responsible for resting the body, while the sympathetic pathway is responsible for preparing for an emergency.
2. Most preganglionic neurons in the sympathetic pathway originate in the spinal cord.
3. Slowing of the heartbeat is a parasympathetic response.
4. Parasympathetic neurons are responsible for releasing norepinephrine on the target organ, while sympathetic neurons are responsible for releasing acetylcholine.
The autonomic nervous system serves as the relay between the CNS and the internal organs. It controls the lungs, the heart, smooth muscle, and exocrine and endocrine glands. The autonomic nervous system controls these organs largely without conscious control; it can continuously monitor the conditions of these different systems and implement changes as needed. Signaling to the target tissue usually involves two synapses: a preganglionic neuron (originating in the CNS) synapses to a neuron in a ganglion that, in turn, synapses on the target organ, as illustrated in Figure \(1\). There are two divisions of the autonomic nervous system that often have opposing effects: the sympathetic nervous system and the parasympathetic nervous system.
Sympathetic Nervous System
The sympathetic nervous system is responsible for the “fight or flight” response that occurs when an animal encounters a dangerous situation. One way to remember this is to think of the surprise a person feels when encountering a snake (“snake” and “sympathetic” both begin with “s”). Examples of functions controlled by the sympathetic nervous system include an accelerated heart rate and inhibited digestion. These functions help prepare an organism’s body for the physical strain required to escape a potentially dangerous situation or to fend off a predator.
Most preganglionic neurons in the sympathetic nervous system originate in the spinal cord, as illustrated in Figure \(2\). The axons of these neurons release acetylcholine on postganglionic neurons within sympathetic ganglia (the sympathetic ganglia form a chain that extends alongside the spinal cord). The acetylcholine activates the postganglionic neurons. Postganglionic neurons then release norepinephrine onto target organs. As anyone who has ever felt a rush before a big test, speech, or athletic event can attest, the effects of the sympathetic nervous system are quite pervasive. This is both because one preganglionic neuron synapses on multiple postganglionic neurons, amplifying the effect of the original synapse, and because the adrenal gland also releases norepinephrine (and the closely related hormone epinephrine) into the blood stream. The physiological effects of this norepinephrine release include dilating the trachea and bronchi (making it easier for the animal to breathe), increasing heart rate, and moving blood from the skin to the heart, muscles, and brain (so the animal can think and run). The strength and speed of the sympathetic response helps an organism avoid danger, and scientists have found evidence that it may also increase LTP—allowing the animal to remember the dangerous situation and avoid it in the future.
Parasympathetic Nervous System
While the sympathetic nervous system is activated in stressful situations, the parasympathetic nervous system allows an animal to “rest and digest.” One way to remember this is to think that during a restful situation like a picnic, the parasympathetic nervous system is in control (“picnic” and “parasympathetic” both start with “p”). Parasympathetic preganglionic neurons have cell bodies located in the brainstem and in the sacral (toward the bottom) spinal cord, as shown in Figure \(2\). The axons of the preganglionic neurons release acetylcholine on the postganglionic neurons, which are generally located very near the target organs. Most postganglionic neurons release acetylcholine onto target organs, although some release nitric oxide.
The parasympathetic nervous system resets organ function after the sympathetic nervous system is activated (the common adrenaline dump you feel after a ‘fight-or-flight’ event). Effects of acetylcholine release on target organs include slowing of heart rate, lowered blood pressure, and stimulation of digestion.
Sensory-Somatic Nervous System
The sensory-somatic nervous system is made up of cranial and spinal nerves and contains both sensory and motor neurons. Sensory neurons transmit sensory information from the skin, skeletal muscle, and sensory organs to the CNS. Motor neurons transmit messages about desired movement from the CNS to the muscles to make them contract. Without its sensory-somatic nervous system, an animal would be unable to process any information about its environment (what it sees, feels, hears, and so on) and could not control motor movements. Unlike the autonomic nervous system, which has two synapses between the CNS and the target organ, sensory and motor neurons have only one synapse—one ending of the neuron is at the organ and the other directly contacts a CNS neuron. Acetylcholine is the main neurotransmitter released at these synapses.
Humans have 12 cranial nerves, nerves that emerge from or enter the skull (cranium), as opposed to the spinal nerves, which emerge from the vertebral column. Each cranial nerve is accorded a name, which are detailed in Figure \(3\). Some cranial nerves transmit only sensory information. For example, the olfactory nerve transmits information about smells from the nose to the brainstem. Other cranial nerves transmit almost solely motor information. For example, the oculomotor nerve controls the opening and closing of the eyelid and some eye movements. Other cranial nerves contain a mix of sensory and motor fibers. For example, the glossopharyngeal nerve has a role in both taste (sensory) and swallowing (motor).
Spinal nerves transmit sensory and motor information between the spinal cord and the rest of the body. Each of the 31 spinal nerves (in humans) contains both sensory and motor axons. The sensory neuron cell bodies are grouped in structures called dorsal root ganglia and are shown in Figure \(4\). Each sensory neuron has one projection—with a sensory receptor ending in skin, muscle, or sensory organs—and another that synapses with a neuron in the dorsal spinal cord. Motor neurons have cell bodies in the ventral gray matter of the spinal cord that project to muscle through the ventral root. These neurons are usually stimulated by interneurons within the spinal cord but are sometimes directly stimulated by sensory neurons.
Summary
The peripheral nervous system contains both the autonomic and sensory-somatic nervous systems. The autonomic nervous system provides unconscious control over visceral functions and has two divisions: the sympathetic and parasympathetic nervous systems. The sympathetic nervous system is activated in stressful situations to prepare the animal for a “fight or flight” response. The parasympathetic nervous system is active during restful periods. The sensory-somatic nervous system is made of cranial and spinal nerves that transmit sensory information from skin and muscle to the CNS and motor commands from the CNS to the muscles.
Art Connections
Figure \(1\): Which of the following statements is false?
1. The parasympathetic pathway is responsible for relaxing the body, while the sympathetic pathway is responsible for preparing for an emergency.
2. Most preganglionic neurons in the sympathetic pathway originate in the spinal cord.
3. Slowing of the heartbeat is a parasympathetic response.
4. Parasympathetic neurons are responsible for releasing norepinephrine on the target organ, while sympathetic neurons are responsible for releasing acetylcholine.
Answer
D
Glossary
acetylcholine
neurotransmitter released by neurons in the central nervous system and peripheral nervous system
autonomic nervous system
part of the peripheral nervous system that controls bodily functions
cranial nerve
sensory and/or motor nerve that emanates from the brain
norepinephrine
neurotransmitter and hormone released by activation of the sympathetic nervous system
parasympathetic nervous system
division of autonomic nervous system that regulates visceral functions during rest and digestion
sensory-somatic nervous system
system of sensory and motor nerves
spinal nerve
nerve projecting between skin or muscle and spinal cord
sympathetic nervous system
division of autonomic nervous system activated during stressful “fight or flight” situations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/42%3A_The_Nervous_System/42.05%3A_The_Peripheral_Nervous_System-_Spinal_and_Cranial_Nerves.txt |
• 43.1: Overview of Sensory Receptors
Senses provide information about the body and its environment. Humans have five special senses: olfaction (smell), gustation (taste), equilibrium (balance and body position), vision, and hearing. Additionally, we possess general senses, also called somatosensation, which respond to stimuli like temperature, pain, pressure, and vibration.
• 43.2: Thermoreceptors- Nociceptors, and Electromagnetic Receptors- Temperature
Somatosensation is a mixed sensory category and includes all sensation received from the skin and mucous membranes, as well from as the limbs and joints. Somatosensation is also known as tactile sense, or more familiarly, as the sense of touch. Somatosensation occurs all over the exterior of the body and at some interior locations as well. A variety of receptor types—embedded in the skin, mucous membranes, muscles, joints, internal organs, and cardiovascular system—play a role.
• 43.3: Mechanoreceptors 1- Touch, Pressure and Body Position
Somatosensation is a mixed sensory category and includes all sensation received from the skin and mucous membranes, as well from as the limbs and joints. Somatosensation is also known as tactile sense, or more familiarly, as the sense of touch. Somatosensation occurs all over the exterior of the body and at some interior locations as well. A variety of receptor types—embedded in the skin, mucous membranes, muscles, joints, internal organs, and cardiovascular system—play a role.
• 43.4: Mechanoreceptors 2- Hearing, Vibration and Balance
• 43.5: Chemoreceptors- Taste, Smell and pH
Taste, also called gustation, and smell, also called olfaction, are the most interconnected senses in that both involve molecules of the stimulus entering the body and bonding to receptors. Smell lets an animal sense the presence of food or other animals—whether potential mates, predators, or prey—or other chemicals in the environment that can impact their survival. Similarly, the sense of taste allows animals to discriminate between types of foods.
• 43.6: Vision
Vision is the ability to detect light patterns from the outside environment and interpret them into images. Animals are bombarded with sensory information, and the sheer volume of visual information can be problematic. Fortunately, the visual systems of species have evolved to attend to the most-important stimuli. The importance of vision to humans is further substantiated by the fact that about one-third of the human cerebral cortex is dedicated to analyzing and perceiving visual information.
• 43.7: Evolution and the Development of Eyes
43: Sensory Systems
Skills to Develop
• Identify the general and special senses in humans
• Describe three important steps in sensory perception
• Explain the concept of just-noticeable difference in sensory perception
Senses provide information about the body and its environment. Humans have five special senses: olfaction (smell), gustation (taste), equilibrium (balance and body position), vision, and hearing. Additionally, we possess general senses, also called somatosensation, which respond to stimuli like temperature, pain, pressure, and vibration. Vestibular sensation, which is an organism’s sense of spatial orientation and balance, proprioception (position of bones, joints, and muscles), and the sense of limb position that is used to track kinesthesia (limb movement) are part of somatosensation. Although the sensory systems associated with these senses are very different, all share a common function: to convert a stimulus (such as light, or sound, or the position of the body) into an electrical signal in the nervous system. This process is called sensory transduction.
There are two broad types of cellular systems that perform sensory transduction. In one, a neuron works with a sensory receptor, a cell, or cell process that is specialized to engage with and detect a specific stimulus. Stimulation of the sensory receptor activates the associated afferent neuron, which carries information about the stimulus to the central nervous system. In the second type of sensory transduction, a sensory nerve ending responds to a stimulus in the internal or external environment: this neuron constitutes the sensory receptor. Free nerve endings can be stimulated by several different stimuli, thus showing little receptor specificity. For example, pain receptors in your gums and teeth may be stimulated by temperature changes, chemical stimulation, or pressure.
Reception
The first step in sensation is reception, which is the activation of sensory receptors by stimuli such as mechanical stimuli (being bent or squished, for example), chemicals, or temperature. The receptor can then respond to the stimuli. The region in space in which a given sensory receptor can respond to a stimulus, be it far away or in contact with the body, is that receptor’s receptive field. Think for a moment about the differences in receptive fields for the different senses. For the sense of touch, a stimulus must come into contact with body. For the sense of hearing, a stimulus can be a moderate distance away (some baleen whale sounds can propagate for many kilometers). For vision, a stimulus can be very far away; for example, the visual system perceives light from stars at enormous distances.
Transduction
The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system. This takes place at the sensory receptor, and the change in electrical potential that is produced is called the receptor potential. How is sensory input, such as pressure on the skin, changed to a receptor potential? In this example, a type of receptor called a mechanoreceptor (as shown in Figure \(1\)) possesses specialized membranes that respond to pressure. Disturbance of these dendrites by compressing them or bending them opens gated ion channels in the plasma membrane of the sensory neuron, changing its electrical potential. Recall that in the nervous system, a positive change of a neuron’s electrical potential (also called the membrane potential), depolarizes the neuron. Receptor potentials are graded potentials: the magnitude of these graded (receptor) potentials varies with the strength of the stimulus. If the magnitude of depolarization is sufficient (that is, if membrane potential reaches a threshold), the neuron will fire an action potential. In most cases, the correct stimulus impinging on a sensory receptor will drive membrane potential in a positive direction, although for some receptors, such as those in the visual system, this is not always the case.
Sensory receptors for different senses are very different from each other, and they are specialized according to the type of stimulus they sense: they have receptor specificity. For example, touch receptors, light receptors, and sound receptors are each activated by different stimuli. Touch receptors are not sensitive to light or sound; they are sensitive only to touch or pressure. However, stimuli may be combined at higher levels in the brain, as happens with olfaction, contributing to our sense of taste.
Encoding and Transmission of Sensory Information
Four aspects of sensory information are encoded by sensory systems: the type of stimulus, the location of the stimulus in the receptive field, the duration of the stimulus, and the relative intensity of the stimulus. Thus, action potentials transmitted over a sensory receptor’s afferent axons encode one type of stimulus, and this segregation of the senses is preserved in other sensory circuits. For example, auditory receptors transmit signals over their own dedicated system, and electrical activity in the axons of the auditory receptors will be interpreted by the brain as an auditory stimulus—a sound.
The intensity of a stimulus is often encoded in the rate of action potentials produced by the sensory receptor. Thus, an intense stimulus will produce a more rapid train of action potentials, and reducing the stimulus will likewise slow the rate of production of action potentials. A second way in which intensity is encoded is by the number of receptors activated. An intense stimulus might initiate action potentials in a large number of adjacent receptors, while a less intense stimulus might stimulate fewer receptors. Integration of sensory information begins as soon as the information is received in the CNS, and the brain will further process incoming signals.
Perception
Perception is an individual’s interpretation of a sensation. Although perception relies on the activation of sensory receptors, perception happens not at the level of the sensory receptor, but at higher levels in the nervous system, in the brain. The brain distinguishes sensory stimuli through a sensory pathway: action potentials from sensory receptors travel along neurons that are dedicated to a particular stimulus. These neurons are dedicated to that particular stimulus and synapse with particular neurons in the brain or spinal cord.
All sensory signals, except those from the olfactory system, are transmitted though the central nervous system and are routed to the thalamus and to the appropriate region of the cortex. Recall that the thalamus is a structure in the forebrain that serves as a clearinghouse and relay station for sensory (as well as motor) signals. When the sensory signal exits the thalamus, it is conducted to the specific area of the cortex (Figure \(2\)) dedicated to processing that particular sense.
How are neural signals interpreted? Interpretation of sensory signals between individuals of the same species is largely similar, owing to the inherited similarity of their nervous systems; however, there are some individual differences. A good example of this is individual tolerances to a painful stimulus, such as dental pain, which certainly differ.
Scientific Method Connection: Just-Noticeable Difference
It is easy to differentiate between a one-pound bag of rice and a two-pound bag of rice. There is a one-pound difference, and one bag is twice as heavy as the other. However, would it be as easy to differentiate between a 20- and a 21-pound bag?
Question: What is the smallest detectible weight difference between a one-pound bag of rice and a larger bag? What is the smallest detectible difference between a 20-pound bag and a larger bag? In both cases, at what weights are the differences detected? This smallest detectible difference in stimuli is known as the just-noticeable difference (JND).
Background: Research background literature on JND and on Weber’s Law, a description of a proposed mathematical relationship between the overall magnitude of the stimulus and the JND. You will be testing JND of different weights of rice in bags. Choose a convenient increment that is to be stepped through while testing. For example, you could choose 10 percent increments between one and two pounds (1.1, 1.2, 1.3, 1.4, and so on) or 20 percent increments (1.2, 1.4, 1.6, and 1.8).
Hypothesis: Develop a hypothesis about JND in terms of percentage of the whole weight being tested (such as “the JND between the two small bags and between the two large bags is proportionally the same,” or “. . . is not proportionally the same.”) So, for the first hypothesis, if the JND between the one-pound bag and a larger bag is 0.2 pounds (that is, 20 percent; 1.0 pound feels the same as 1.1 pounds, but 1.0 pound feels less than 1.2 pounds), then the JND between the 20-pound bag and a larger bag will also be 20 percent. (So, 20 pounds feels the same as 22 pounds or 23 pounds, but 20 pounds feels less than 24 pounds.)
Test the hypothesis: Enlist 24 participants, and split them into two groups of 12. To set up the demonstration, assuming a 10 percent increment was selected, have the first group be the one-pound group. As a counter-balancing measure against a systematic error, however, six of the first group will compare one pound to two pounds, and step down in weight (1.0 to 2.0, 1.0 to 1.9, and so on.), while the other six will step up (1.0 to 1.1, 1.0 to 1.2, and so on). Apply the same principle to the 20-pound group (20 to 40, 20 to 38, and so on, and 20 to 22, 20 to 24, and so on). Given the large difference between 20 and 40 pounds, you may wish to use 30 pounds as your larger weight. In any case, use two weights that are easily detectable as different.
Record the observations: Record the data in a table similar to the table below. For the one-pound and 20-pound groups (base weights) record a plus sign (+) for each participant that detects a difference between the base weight and the step weight. Record a minus sign (-) for each participant that finds no difference. If one-tenth steps were not used, then replace the steps in the “Step Weight” columns with the step you are using.
Table \(1\): Results of JND Testing (+ = difference; – = no difference)
Step Weight One pound 20 pounds Step Weight
1.1 22
1.2 24
1.3 26
1.4 28
1.5 30
1.6 32
1.7 34
1.8 36
1.9 38
2.0 40
Analyze the data/report the results: What step weight did all participants find to be equal with one-pound base weight? What about the 20-pound group?
Draw a conclusion: Did the data support the hypothesis? Are the final weights proportionally the same? If not, why not? Do the findings adhere to Weber’s Law? Weber’s Law states that the concept that a just-noticeable difference in a stimulus is proportional to the magnitude of the original stimulus.
Summary
A sensory activation occurs when a physical or chemical stimulus is processed into a neural signal (sensory transduction) by a sensory receptor. Perception is an individual interpretation of a sensation and is a brain function. Humans have special senses: olfaction, gustation, equilibrium, and hearing, plus the general senses of somatosensation.
Sensory receptors are either specialized cells associated with sensory neurons or the specialized ends of sensory neurons that are a part of the peripheral nervous system, and they are used to receive information about the environment (internal or external). Each sensory receptor is modified for the type of stimulus it detects. For example, neither gustatory receptors nor auditory receptors are sensitive to light. Each sensory receptor is responsive to stimuli within a specific region in space, which is known as that receptor’s receptive field. The most fundamental function of a sensory system is the translation of a sensory signal to an electrical signal in the nervous system.
All sensory signals, except those from the olfactory system, enter the central nervous system and are routed to the thalamus. When the sensory signal exits the thalamus, it is conducted to the specific area of the cortex dedicated to processing that particular sense.
Glossary
kinesthesia
sense of body movement
mechanoreceptor
sensory receptor modified to respond to mechanical disturbance such as being bent, touch, pressure, motion, and sound
perception
individual interpretation of a sensation; a brain function
proprioception
sense of limb position; used to track kinesthesia
reception
receipt of a signal (such as light or sound) by sensory receptors
receptive field
region in space in which a stimulus can activate a given sensory receptor
receptor potential
membrane potential in a sensory receptor in response to detection of a stimulus
sensory receptor
specialized neuron or other cells associated with a neuron that is modified to receive specific sensory input
sensory transduction
conversion of a sensory stimulus into electrical energy in the nervous system by a change in the membrane potential
vestibular sense
sense of spatial orientation and balance | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/43%3A_Sensory_Systems/43.01%3A_Overview_of_Sensory_Receptors.txt |
Skills to Develop
• Describe four important mechanoreceptors in human skin
• Describe the topographical distribution of somatosensory receptors between glabrous and hairy skin
• Explain why the perception of pain is subjective
Somatosensation is a mixed sensory category and includes all sensation received from the skin and mucous membranes, as well from as the limbs and joints. Somatosensation is also known as tactile sense, or more familiarly, as the sense of touch. Somatosensation occurs all over the exterior of the body and at some interior locations as well. A variety of receptor types—embedded in the skin, mucous membranes, muscles, joints, internal organs, and cardiovascular system—play a role.
Recall that the epidermis is the outermost layer of skin in mammals. It is relatively thin, is composed of keratin-filled cells, and has no blood supply. The epidermis serves as a barrier to water and to invasion by pathogens. Below this, the much thicker dermis contains blood vessels, sweat glands, hair follicles, lymph vessels, and lipid-secreting sebaceous glands (Figure \(1\)). Below the epidermis and dermis is the subcutaneous tissue, or hypodermis, the fatty layer that contains blood vessels, connective tissue, and the axons of sensory neurons. The hypodermis, which holds about 50 percent of the body’s fat, attaches the dermis to the bone and muscle, and supplies nerves and blood vessels to the dermis.
Somatosensory Receptors
Sensory receptors are classified into five categories: mechanoreceptors, thermoreceptors, proprioceptors, pain receptors, and chemoreceptors. These categories are based on the nature of stimuli each receptor class transduces. What is commonly referred to as “touch” involves more than one kind of stimulus and more than one kind of receptor. Mechanoreceptors in the skin are described as encapsulated (that is, surrounded by a capsule) or unencapsulated (a group that includes free nerve endings). A free nerve ending, as its name implies, is an unencapsulated dendrite of a sensory neuron. Free nerve endings are the most common nerve endings in skin, and they extend into the middle of the epidermis. Free nerve endings are sensitive to painful stimuli, to hot and cold, and to light touch. They are slow to adjust to a stimulus and so are less sensitive to abrupt changes in stimulation.
There are three classes of mechanoreceptors: tactile, proprioceptors, and baroreceptors. Mechanoreceptors sense stimuli due to physical deformation of their plasma membranes. They contain mechanically gated ion channels whose gates open or close in response to pressure, touch, stretching, and sound.” There are four primary tactile mechanoreceptors in human skin: Merkel’s disks, Meissner’s corpuscles, Ruffini endings, and Pacinian corpuscle; two are located toward the surface of the skin and two are located deeper. A fifth type of mechanoreceptor, Krause end bulbs, are found only in specialized regions. Merkel’s disks (shown in Figure \(2\)) are found in the upper layers of skin near the base of the epidermis, both in skin that has hair and on glabrous skin, that is, the hairless skin found on the palms and fingers, the soles of the feet, and the lips of humans and other primates. Merkel’s disks are densely distributed in the fingertips and lips. They are slow-adapting, unencapsulated nerve endings, and they respond to light touch. Light touch, also known as discriminative touch, is a light pressure that allows the location of a stimulus to be pinpointed. The receptive fields of Merkel’s disks are small with well-defined borders. That makes them finely sensitive to edges and they come into use in tasks such as typing on a keyboard.
Exercise
Which of the following statements about mechanoreceptors is false?
1. Pacini corpuscles are found in both glabrous and hairy skin.
2. Merkel’s disks are abundant on the fingertips and lips.
3. Ruffini endings are encapsulated mechanoreceptors.
4. Meissner’s corpuscles extend into the lower dermis.
Answer
D
Meissner’s corpuscles, (shown in Figure \(3\)) also known as tactile corpuscles, are found in the upper dermis, but they project into the epidermis. They, too, are found primarily in the glabrous skin on the fingertips and eyelids. They respond to fine touch and pressure, but they also respond to low-frequency vibration or flutter. They are rapidly adapting, fluid-filled, encapsulated neurons with small, well-defined borders and are responsive to fine details. Like Merkel’s disks, Meissner’s corpuscles are not as plentiful in the palms as they are in the fingertips.
Deeper in the epidermis, near the base, are Ruffini endings, which are also known as bulbous corpuscles. They are found in both glabrous and hairy skin. These are slow-adapting, encapsulated mechanoreceptors that detect skin stretch and deformations within joints, so they provide valuable feedback for gripping objects and controlling finger position and movement. Thus, they also contribute to proprioception and kinesthesia. Ruffini endings also detect warmth. Note that these warmth detectors are situated deeper in the skin than are the cold detectors. It is not surprising, then, that humans detect cold stimuli before they detect warm stimuli.
Pacinian corpuscles (seen in Figure \(4\)) are located deep in the dermis of both glabrous and hairy skin and are structurally similar to Meissner’s corpuscles; they are found in the bone periosteum, joint capsules, pancreas and other viscera, breast, and genitals. They are rapidly adapting mechanoreceptors that sense deep transient (but not prolonged) pressure and high-frequency vibration. Pacinian receptors detect pressure and vibration by being compressed, stimulating their internal dendrites. There are fewer Pacinian corpuscles and Ruffini endings in skin than there are Merkel’s disks and Meissner’s corpuscles.
In proprioception, proprioceptive and kinesthetic signals travel through myelinated afferent neurons running from the spinal cord to the medulla. Neurons are not physically connected, but communicate via neurotransmitters secreted into synapses or “gaps” between communicating neurons. Once in the medulla, the neurons continue carrying the signals to the thalamus.
Muscle spindles are stretch receptors that detect the amount of stretch, or lengthening of muscles. Related to these are Golgi tendon organs, which are tension receptors that detect the force of muscle contraction. Proprioceptive and kinesthetic signals come from limbs. Unconscious proprioceptive signals run from the spinal cord to the cerebellum, the brain region that coordinates muscle contraction, rather than to the thalamus, like most other sensory information.
Barorecptors detect pressure changes in an organ. They are found in the walls of the carotid artery and the aorta where they monitor blood pressure, and in the lungs where they detect the degree of lung expansion. Stretch receptors are found at various sites in the digestive and urinary systems.
In addition to these two types of deeper receptors, there are also rapidly adapting hair receptors, which are found on nerve endings that wrap around the base of hair follicles. There are a few types of hair receptors that detect slow and rapid hair movement, and they differ in their sensitivity to movement. Some hair receptors also detect skin deflection, and certain rapidly adapting hair receptors allow detection of stimuli that have not yet touched the skin.
Integration of Signals from Mechanoreceptors
The configuration of the different types of receptors working in concert in human skin results in a very refined sense of touch. The nociceptive receptors—those that detect pain—are located near the surface. Small, finely calibrated mechanoreceptors—Merkel’s disks and Meissner’s corpuscles—are located in the upper layers and can precisely localize even gentle touch. The large mechanoreceptors—Pacinian corpuscles and Ruffini endings—are located in the lower layers and respond to deeper touch. (Consider that the deep pressure that reaches those deeper receptors would not need to be finely localized.) Both the upper and lower layers of the skin hold rapidly and slowly adapting receptors. Both primary somatosensory cortex and secondary cortical areas are responsible for processing the complex picture of stimuli transmitted from the interplay of mechanoreceptors.
Density of Mechanoreceptors
The distribution of touch receptors in human skin is not consistent over the body. In humans, touch receptors are less dense in skin covered with any type of hair, such as the arms, legs, torso, and face. Touch receptors are denser in glabrous skin (the type found on human fingertips and lips, for example), which is typically more sensitive and is thicker than hairy skin (4 to 5 mm versus 2 to 3 mm).
How is receptor density estimated in a human subject? The relative density of pressure receptors in different locations on the body can be demonstrated experimentally using a two-point discrimination test. In this demonstration, two sharp points, such as two thumbtacks, are brought into contact with the subject’s skin (though not hard enough to cause pain or break the skin). The subject reports if he or she feels one point or two points. If the two points are felt as one point, it can be inferred that the two points are both in the receptive field of a single sensory receptor. If two points are felt as two separate points, each is in the receptive field of two separate sensory receptors. The points could then be moved closer and re-tested until the subject reports feeling only one point, and the size of the receptive field of a single receptor could be estimated from that distance.
Thermoreception
In addition to Krause end bulbs that detect cold and Ruffini endings that detect warmth, there are different types of cold receptors on some free nerve endings: thermoreceptors, located in the dermis, skeletal muscles, liver, and hypothalamus, that are activated by different temperatures. Their pathways into the brain run from the spinal cord through the thalamus to the primary somatosensory cortex. Warmth and cold information from the face travels through one of the cranial nerves to the brain. You know from experience that a tolerably cold or hot stimulus can quickly progress to a much more intense stimulus that is no longer tolerable. Any stimulus that is too intense can be perceived as pain because temperature sensations are conducted along the same pathways that carry pain sensations
Pain
Pain is the name given to nociception, which is the neural processing of injurious stimuli in response to tissue damage. Pain is caused by true sources of injury, such as contact with a heat source that causes a thermal burn or contact with a corrosive chemical. But pain also can be caused by harmless stimuli that mimic the action of damaging stimuli, such as contact with capsaicins, the compounds that cause peppers to taste hot and which are used in self-defense pepper sprays and certain topical medications. Peppers taste “hot” because the protein receptors that bind capsaicin open the same calcium channels that are activated by warm receptors.
Nociception starts at the sensory receptors, but pain, inasmuch as it is the perception of nociception, does not start until it is communicated to the brain. There are several nociceptive pathways to and through the brain. Most axons carrying nociceptive information into the brain from the spinal cord project to the thalamus (as do other sensory neurons) and the neural signal undergoes final processing in the primary somatosensory cortex. Interestingly, one nociceptive pathway projects not to the thalamus but directly to the hypothalamus in the forebrain, which modulates the cardiovascular and neuroendocrine functions of the autonomic nervous system. Recall that threatening—or painful—stimuli stimulate the sympathetic branch of the visceral sensory system, readying a fight-or-flight response.
Link to Learning
View this video that animates the five phases of nociceptive pain.
Summary
Somatosensation includes all sensation received from the skin and mucous membranes, as well as from the limbs and joints. Somatosensation occurs all over the exterior of the body and at some interior locations as well, and a variety of receptor types, embedded in the skin and mucous membranes, play a role.
There are several types of specialized sensory receptors. Rapidly adapting free nerve endings detect nociception, hot and cold, and light touch. Slowly adapting, encapsulated Merkel’s disks are found in fingertips and lips, and respond to light touch. Meissner’s corpuscles, found in glabrous skin, are rapidly adapting, encapsulated receptors that detect touch, low-frequency vibration, and flutter. Ruffini endings are slowly adapting, encapsulated receptors that detect skin stretch, joint activity, and warmth. Hair receptors are rapidly adapting nerve endings wrapped around the base of hair follicles that detect hair movement and skin deflection. Finally, Pacinian corpuscles are encapsulated, rapidly adapting receptors that detect transient pressure and high-frequency vibration.
Glossary
free nerve ending
ending of an afferent neuron that lacks a specialized structure for detection of sensory stimuli; some respond to touch, pain, or temperature
glabrous
describes the non-hairy skin found on palms and fingers, soles of feet, and lips of humans and other primates
Golgi tendon organ
muscular proprioceptive tension receptor that provides the sensory component of the Golgi tendon reflex
Meissner’s corpuscle
(also, tactile corpuscle) encapsulated, rapidly-adapting mechanoreceptor in the skin that responds to light touch
Merkel's disc
unencapsulated, slowly-adapting mechanoreceptor in the skin that responds to touch
muscle spindle
proprioceptive stretch receptor that lies within a muscle and that shortens the muscle to an optimal length for efficient contraction
nociception
neural processing of noxious (such as damaging) stimuli
Pacinian corpuscle
encapsulated mechanoreceptor in the skin that responds to deep pressure and vibration
Ruffini ending
(also, bulbous corpuscle) slowly-adapting mechanoreceptor in the skin that responds to skin stretch and joint position | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/43%3A_Sensory_Systems/43.02%3A_Thermoreceptors-_Nociceptors_and_Electromagnetic_Receptors-_Temperature.txt |
Skills to Develop
• Describe four important mechanoreceptors in human skin
• Describe the topographical distribution of somatosensory receptors between glabrous and hairy skin
• Explain why the perception of pain is subjective
Somatosensation is a mixed sensory category and includes all sensation received from the skin and mucous membranes, as well from as the limbs and joints. Somatosensation is also known as tactile sense, or more familiarly, as the sense of touch. Somatosensation occurs all over the exterior of the body and at some interior locations as well. A variety of receptor types—embedded in the skin, mucous membranes, muscles, joints, internal organs, and cardiovascular system—play a role.
Recall that the epidermis is the outermost layer of skin in mammals. It is relatively thin, is composed of keratin-filled cells, and has no blood supply. The epidermis serves as a barrier to water and to invasion by pathogens. Below this, the much thicker dermis contains blood vessels, sweat glands, hair follicles, lymph vessels, and lipid-secreting sebaceous glands (Figure \(1\)). Below the epidermis and dermis is the subcutaneous tissue, or hypodermis, the fatty layer that contains blood vessels, connective tissue, and the axons of sensory neurons. The hypodermis, which holds about 50 percent of the body’s fat, attaches the dermis to the bone and muscle, and supplies nerves and blood vessels to the dermis.
Somatosensory Receptors
Sensory receptors are classified into five categories: mechanoreceptors, thermoreceptors, proprioceptors, pain receptors, and chemoreceptors. These categories are based on the nature of stimuli each receptor class transduces. What is commonly referred to as “touch” involves more than one kind of stimulus and more than one kind of receptor. Mechanoreceptors in the skin are described as encapsulated (that is, surrounded by a capsule) or unencapsulated (a group that includes free nerve endings). A free nerve ending, as its name implies, is an unencapsulated dendrite of a sensory neuron. Free nerve endings are the most common nerve endings in skin, and they extend into the middle of the epidermis. Free nerve endings are sensitive to painful stimuli, to hot and cold, and to light touch. They are slow to adjust to a stimulus and so are less sensitive to abrupt changes in stimulation.
There are three classes of mechanoreceptors: tactile, proprioceptors, and baroreceptors. Mechanoreceptors sense stimuli due to physical deformation of their plasma membranes. They contain mechanically gated ion channels whose gates open or close in response to pressure, touch, stretching, and sound.” There are four primary tactile mechanoreceptors in human skin: Merkel’s disks, Meissner’s corpuscles, Ruffini endings, and Pacinian corpuscle; two are located toward the surface of the skin and two are located deeper. A fifth type of mechanoreceptor, Krause end bulbs, are found only in specialized regions. Merkel’s disks (shown in Figure \(2\)) are found in the upper layers of skin near the base of the epidermis, both in skin that has hair and on glabrous skin, that is, the hairless skin found on the palms and fingers, the soles of the feet, and the lips of humans and other primates. Merkel’s disks are densely distributed in the fingertips and lips. They are slow-adapting, unencapsulated nerve endings, and they respond to light touch. Light touch, also known as discriminative touch, is a light pressure that allows the location of a stimulus to be pinpointed. The receptive fields of Merkel’s disks are small with well-defined borders. That makes them finely sensitive to edges and they come into use in tasks such as typing on a keyboard.
Exercise
Which of the following statements about mechanoreceptors is false?
1. Pacini corpuscles are found in both glabrous and hairy skin.
2. Merkel’s disks are abundant on the fingertips and lips.
3. Ruffini endings are encapsulated mechanoreceptors.
4. Meissner’s corpuscles extend into the lower dermis.
Answer
D
Meissner’s corpuscles, (shown in Figure \(3\)) also known as tactile corpuscles, are found in the upper dermis, but they project into the epidermis. They, too, are found primarily in the glabrous skin on the fingertips and eyelids. They respond to fine touch and pressure, but they also respond to low-frequency vibration or flutter. They are rapidly adapting, fluid-filled, encapsulated neurons with small, well-defined borders and are responsive to fine details. Like Merkel’s disks, Meissner’s corpuscles are not as plentiful in the palms as they are in the fingertips.
Deeper in the epidermis, near the base, are Ruffini endings, which are also known as bulbous corpuscles. They are found in both glabrous and hairy skin. These are slow-adapting, encapsulated mechanoreceptors that detect skin stretch and deformations within joints, so they provide valuable feedback for gripping objects and controlling finger position and movement. Thus, they also contribute to proprioception and kinesthesia. Ruffini endings also detect warmth. Note that these warmth detectors are situated deeper in the skin than are the cold detectors. It is not surprising, then, that humans detect cold stimuli before they detect warm stimuli.
Pacinian corpuscles (seen in Figure \(4\)) are located deep in the dermis of both glabrous and hairy skin and are structurally similar to Meissner’s corpuscles; they are found in the bone periosteum, joint capsules, pancreas and other viscera, breast, and genitals. They are rapidly adapting mechanoreceptors that sense deep transient (but not prolonged) pressure and high-frequency vibration. Pacinian receptors detect pressure and vibration by being compressed, stimulating their internal dendrites. There are fewer Pacinian corpuscles and Ruffini endings in skin than there are Merkel’s disks and Meissner’s corpuscles.
In proprioception, proprioceptive and kinesthetic signals travel through myelinated afferent neurons running from the spinal cord to the medulla. Neurons are not physically connected, but communicate via neurotransmitters secreted into synapses or “gaps” between communicating neurons. Once in the medulla, the neurons continue carrying the signals to the thalamus.
Muscle spindles are stretch receptors that detect the amount of stretch, or lengthening of muscles. Related to these are Golgi tendon organs, which are tension receptors that detect the force of muscle contraction. Proprioceptive and kinesthetic signals come from limbs. Unconscious proprioceptive signals run from the spinal cord to the cerebellum, the brain region that coordinates muscle contraction, rather than to the thalamus, like most other sensory information.
Barorecptors detect pressure changes in an organ. They are found in the walls of the carotid artery and the aorta where they monitor blood pressure, and in the lungs where they detect the degree of lung expansion. Stretch receptors are found at various sites in the digestive and urinary systems.
In addition to these two types of deeper receptors, there are also rapidly adapting hair receptors, which are found on nerve endings that wrap around the base of hair follicles. There are a few types of hair receptors that detect slow and rapid hair movement, and they differ in their sensitivity to movement. Some hair receptors also detect skin deflection, and certain rapidly adapting hair receptors allow detection of stimuli that have not yet touched the skin.
Integration of Signals from Mechanoreceptors
The configuration of the different types of receptors working in concert in human skin results in a very refined sense of touch. The nociceptive receptors—those that detect pain—are located near the surface. Small, finely calibrated mechanoreceptors—Merkel’s disks and Meissner’s corpuscles—are located in the upper layers and can precisely localize even gentle touch. The large mechanoreceptors—Pacinian corpuscles and Ruffini endings—are located in the lower layers and respond to deeper touch. (Consider that the deep pressure that reaches those deeper receptors would not need to be finely localized.) Both the upper and lower layers of the skin hold rapidly and slowly adapting receptors. Both primary somatosensory cortex and secondary cortical areas are responsible for processing the complex picture of stimuli transmitted from the interplay of mechanoreceptors.
Density of Mechanoreceptors
The distribution of touch receptors in human skin is not consistent over the body. In humans, touch receptors are less dense in skin covered with any type of hair, such as the arms, legs, torso, and face. Touch receptors are denser in glabrous skin (the type found on human fingertips and lips, for example), which is typically more sensitive and is thicker than hairy skin (4 to 5 mm versus 2 to 3 mm).
How is receptor density estimated in a human subject? The relative density of pressure receptors in different locations on the body can be demonstrated experimentally using a two-point discrimination test. In this demonstration, two sharp points, such as two thumbtacks, are brought into contact with the subject’s skin (though not hard enough to cause pain or break the skin). The subject reports if he or she feels one point or two points. If the two points are felt as one point, it can be inferred that the two points are both in the receptive field of a single sensory receptor. If two points are felt as two separate points, each is in the receptive field of two separate sensory receptors. The points could then be moved closer and re-tested until the subject reports feeling only one point, and the size of the receptive field of a single receptor could be estimated from that distance.
Thermoreception
In addition to Krause end bulbs that detect cold and Ruffini endings that detect warmth, there are different types of cold receptors on some free nerve endings: thermoreceptors, located in the dermis, skeletal muscles, liver, and hypothalamus, that are activated by different temperatures. Their pathways into the brain run from the spinal cord through the thalamus to the primary somatosensory cortex. Warmth and cold information from the face travels through one of the cranial nerves to the brain. You know from experience that a tolerably cold or hot stimulus can quickly progress to a much more intense stimulus that is no longer tolerable. Any stimulus that is too intense can be perceived as pain because temperature sensations are conducted along the same pathways that carry pain sensations
Pain
Pain is the name given to nociception, which is the neural processing of injurious stimuli in response to tissue damage. Pain is caused by true sources of injury, such as contact with a heat source that causes a thermal burn or contact with a corrosive chemical. But pain also can be caused by harmless stimuli that mimic the action of damaging stimuli, such as contact with capsaicins, the compounds that cause peppers to taste hot and which are used in self-defense pepper sprays and certain topical medications. Peppers taste “hot” because the protein receptors that bind capsaicin open the same calcium channels that are activated by warm receptors.
Nociception starts at the sensory receptors, but pain, inasmuch as it is the perception of nociception, does not start until it is communicated to the brain. There are several nociceptive pathways to and through the brain. Most axons carrying nociceptive information into the brain from the spinal cord project to the thalamus (as do other sensory neurons) and the neural signal undergoes final processing in the primary somatosensory cortex. Interestingly, one nociceptive pathway projects not to the thalamus but directly to the hypothalamus in the forebrain, which modulates the cardiovascular and neuroendocrine functions of the autonomic nervous system. Recall that threatening—or painful—stimuli stimulate the sympathetic branch of the visceral sensory system, readying a fight-or-flight response.
Link to Learning
View this video that animates the five phases of nociceptive pain.
Summary
Somatosensation includes all sensation received from the skin and mucous membranes, as well as from the limbs and joints. Somatosensation occurs all over the exterior of the body and at some interior locations as well, and a variety of receptor types, embedded in the skin and mucous membranes, play a role.
There are several types of specialized sensory receptors. Rapidly adapting free nerve endings detect nociception, hot and cold, and light touch. Slowly adapting, encapsulated Merkel’s disks are found in fingertips and lips, and respond to light touch. Meissner’s corpuscles, found in glabrous skin, are rapidly adapting, encapsulated receptors that detect touch, low-frequency vibration, and flutter. Ruffini endings are slowly adapting, encapsulated receptors that detect skin stretch, joint activity, and warmth. Hair receptors are rapidly adapting nerve endings wrapped around the base of hair follicles that detect hair movement and skin deflection. Finally, Pacinian corpuscles are encapsulated, rapidly adapting receptors that detect transient pressure and high-frequency vibration.
Glossary
free nerve ending
ending of an afferent neuron that lacks a specialized structure for detection of sensory stimuli; some respond to touch, pain, or temperature
glabrous
describes the non-hairy skin found on palms and fingers, soles of feet, and lips of humans and other primates
Golgi tendon organ
muscular proprioceptive tension receptor that provides the sensory component of the Golgi tendon reflex
Meissner’s corpuscle
(also, tactile corpuscle) encapsulated, rapidly-adapting mechanoreceptor in the skin that responds to light touch
Merkel's disc
unencapsulated, slowly-adapting mechanoreceptor in the skin that responds to touch
muscle spindle
proprioceptive stretch receptor that lies within a muscle and that shortens the muscle to an optimal length for efficient contraction
nociception
neural processing of noxious (such as damaging) stimuli
Pacinian corpuscle
encapsulated mechanoreceptor in the skin that responds to deep pressure and vibration
Ruffini ending
(also, bulbous corpuscle) slowly-adapting mechanoreceptor in the skin that responds to skin stretch and joint position | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/43%3A_Sensory_Systems/43.03%3A_Mechanoreceptors_1-_Touch_Pressure_and_Body_Position.txt |
Skills to Develop
• Describe the relationship of amplitude and frequency of a sound wave to attributes of sound
• Trace the path of sound through the auditory system to the site of transduction of sound
• Identify the structures of the vestibular system that respond to gravity
Audition, or hearing, is important to humans and to other animals for many different interactions. It enables an organism to detect and receive information about danger, such as an approaching predator, and to participate in communal exchanges like those concerning territories or mating. On the other hand, although it is physically linked to the auditory system, the vestibular system is not involved in hearing. Instead, an animal’s vestibular system detects its own movement, both linear and angular acceleration and deceleration, and balance.
Sound
Auditory stimuli are sound waves, which are mechanical, pressure waves that move through a medium, such as air or water. There are no sound waves in a vacuum since there are no air molecules to move in waves. The speed of sound waves differs, based on altitude, temperature, and medium, but at sea level and a temperature of 20º C (68º F), sound waves travel in the air at about 343 meters per second.
As is true for all waves, there are four main characteristics of a sound wave: frequency, wavelength, period, and amplitude. Frequency is the number of waves per unit of time, and in sound is heard as pitch. High-frequency (≥15.000Hz) sounds are higher-pitched (short wavelength) than low-frequency (long wavelengths; ≤100Hz) sounds. Frequency is measured in cycles per second, and for sound, the most commonly used unit is hertz (Hz), or cycles per second. Most humans can perceive sounds with frequencies between 30 and 20,000 Hz. Women are typically better at hearing high frequencies, but everyone’s ability to hear high frequencies decreases with age. Dogs detect up to about 40,000 Hz; cats, 60,000 Hz; bats, 100,000 Hz; and dolphins 150,000 Hz, and American shad (Alosa sapidissima), a fish, can hear 180,000 Hz. Those frequencies above the human range are called ultrasound.
Amplitude, or the dimension of a wave from peak to trough, in sound is heard as volume and is illustrated in Figure \(1\). The sound waves of louder sounds have greater amplitude than those of softer sounds. For sound, volume is measured in decibels (dB). The softest sound that a human can hear is the zero point. Humans speak normally at 60 decibels.
Reception of Sound
In mammals, sound waves are collected by the external, cartilaginous part of the ear called the pinna, then travel through the auditory canal and cause vibration of the thin diaphragm called the tympanum or ear drum, the innermost part of the outer ear (illustrated in Figure \(2\)). Interior to the tympanum is the middle ear. The middle ear holds three small bones called the ossicles, which transfer energy from the moving tympanum to the inner ear. The three ossicles are the malleus (also known as the hammer), the incus (the anvil), and stapes (the stirrup). The aptly named stapes looks very much like a stirrup. The three ossicles are unique to mammals, and each plays a role in hearing. The malleus attaches at three points to the interior surface of the tympanic membrane. The incus attaches the malleus to the stapes. In humans, the stapes is not long enough to reach the tympanum. If we did not have the malleus and the incus, then the vibrations of the tympanum would never reach the inner ear. These bones also function to collect force and amplify sounds. The ear ossicles are homologous to bones in a fish mouth: the bones that support gills in fish are thought to be adapted for use in the vertebrate ear over evolutionary time. Many animals (frogs, reptiles, and birds, for example) use the stapes of the middle ear to transmit vibrations to the middle ear.
Transduction of Sound
Vibrating objects, such as vocal cords, create sound waves or pressure waves in the air. When these pressure waves reach the ear, the ear transduces this mechanical stimulus (pressure wave) into a nerve impulse (electrical signal) that the brain perceives as sound. The pressure waves strike the tympanum, causing it to vibrate. The mechanical energy from the moving tympanum transmits the vibrations to the three bones of the middle ear. The stapes transmits the vibrations to a thin diaphragm called the oval window, which is the outermost structure of the inner ear. The structures of the inner ear are found in the labyrinth, a bony, hollow structure that is the most interior portion of the ear. Here, the energy from the sound wave is transferred from the stapes through the flexible oval window and to the fluid of the cochlea. The vibrations of the oval window create pressure waves in the fluid (perilymph) inside the cochlea. The cochlea is a whorled structure, like the shell of a snail, and it contains receptors for transduction of the mechanical wave into an electrical signal (as illustrated in Figure \(3\)). Inside the cochlea, the basilar membrane is a mechanical analyzer that runs the length of the cochlea, curling toward the cochlea’s center.
The mechanical properties of the basilar membrane change along its length, such that it is thicker, tauter, and narrower at the outside of the whorl (where the cochlea is largest), and thinner, floppier, and broader toward the apex, or center, of the whorl (where the cochlea is smallest). Different regions of the basilar membrane vibrate according to the frequency of the sound wave conducted through the fluid in the cochlea. For these reasons, the fluid-filled cochlea detects different wave frequencies (pitches) at different regions of the membrane. When the sound waves in the cochlear fluid contact the basilar membrane, it flexes back and forth in a wave-like fashion. Above the basilar membrane is the tectorial membrane.
Exercise
Cochlear implants can restore hearing in people who have a nonfunctional cochlear. The implant consists of a microphone that picks up sound. A speech processor selects sounds in the range of human speech, and a transmitter converts these sounds to electrical impulses, which are then sent to the auditory nerve. Which of the following types of hearing loss would not be restored by a cochlear implant?
1. Hearing loss resulting from absence or loss of hair cells in the organ of Corti.
2. Hearing loss resulting from an abnormal auditory nerve.
3. Hearing loss resulting from fracture of the cochlea.
4. Hearing loss resulting from damage to bones of the middle ear.
Answer
B
The site of transduction is in the organ of Corti (spiral organ). It is composed of hair cells held in place above the basilar membrane like flowers projecting up from soil, with their exposed short, hair-like stereocilia contacting or embedded in the tectorial membrane above them. The inner hair cells are the primary auditory receptors and exist in a single row, numbering approximately 3,500. The stereocilia from inner hair cells extend into small dimples on the tectorial membrane’s lower surface. The outer hair cells are arranged in three or four rows. They number approximately 12,000, and they function to fine tune incoming sound waves. The longer stereocilia that project from the outer hair cells actually attach to the tectorial membrane. All of the stereocilia are mechanoreceptors, and when bent by vibrations they respond by opening a gated ion channel. As a result, the hair cell membrane is depolarized, and a signal is transmitted to the chochlear nerve. Intensity (volume) of sound is determined by how many hair cells at a particular location are stimulated.
The hair cells are arranged on the basilar membrane in an orderly way. The basilar membrane vibrates in different regions, according to the frequency of the sound waves impinging on it. Likewise, the hair cells that lay above it are most sensitive to a specific frequency of sound waves. Hair cells can respond to a small range of similar frequencies, but they require stimulation of greater intensity to fire at frequencies outside of their optimal range. The difference in response frequency between adjacent inner hair cells is about 0.2 percent. Compare that to adjacent piano strings, which are about six percent different. Place theory, which is the model for how biologists think pitch detection works in the human ear, states that high frequency sounds selectively vibrate the basilar membrane of the inner ear near the entrance port (the oval window). Lower frequencies travel farther along the membrane before causing appreciable excitation of the membrane. The basic pitch-determining mechanism is based on the location along the membrane where the hair cells are stimulated. The place theory is the first step toward an understanding of pitch perception. Considering the extreme pitch sensitivity of the human ear, it is thought that there must be some auditory “sharpening” mechanism to enhance the pitch resolution.
When sound waves produce fluid waves inside the cochlea, the basilar membrane flexes, bending the stereocilia that attach to the tectorial membrane. Their bending results in action potentials in the hair cells, and auditory information travels along the neural endings of the bipolar neurons of the hair cells (collectively, the auditory nerve) to the brain. When the hairs bend, they release an excitatory neurotransmitter at a synapse with a sensory neuron, which then conducts action potentials to the central nervous system. The cochlear branch of the vestibulocochlear cranial nerve sends information on hearing. The auditory system is very refined, and there is some modulation or “sharpening” built in. The brain can send signals back to the cochlea, resulting in a change of length in the outer hair cells, sharpening or dampening the hair cells’ response to certain frequencies.
Higher Processing
The inner hair cells are most important for conveying auditory information to the brain. About 90 percent of the afferent neurons carry information from inner hair cells, with each hair cell synapsing with 10 or so neurons. Outer hair cells connect to only 10 percent of the afferent neurons, and each afferent neuron innervates many hair cells. The afferent, bipolar neurons that convey auditory information travel from the cochlea to the medulla, through the pons and midbrain in the brainstem, finally reaching the primary auditory cortex in the temporal lobe.
Vestibular Information
The stimuli associated with the vestibular system are linear acceleration (gravity) and angular acceleration and deceleration. Gravity, acceleration, and deceleration are detected by evaluating the inertia on receptive cells in the vestibular system. Gravity is detected through head position. Angular acceleration and deceleration are expressed through turning or tilting of the head.
The vestibular system has some similarities with the auditory system. It utilizes hair cells just like the auditory system, but it excites them in different ways. There are five vestibular receptor organs in the inner ear: the utricle, the saccule, and three semicircular canals. Together, they make up what’s known as the vestibular labyrinth that is shown in Figure \(4\). The utricle and saccule respond to acceleration in a straight line, such as gravity. The roughly 30,000 hair cells in the utricle and 16,000 hair cells in the saccule lie below a gelatinous layer, with their stereocilia projecting into the gelatin. Embedded in this gelatin are calcium carbonate crystals—like tiny rocks. When the head is tilted, the crystals continue to be pulled straight down by gravity, but the new angle of the head causes the gelatin to shift, thereby bending the stereocilia. The bending of the stereocilia stimulates the neurons, and they signal to the brain that the head is tilted, allowing the maintenance of balance. It is the vestibular branch of the vestibulocochlear cranial nerve that deals with balance.
The fluid-filled semicircular canals are tubular loops set at oblique angles. They are arranged in three spatial planes. The base of each canal has a swelling that contains a cluster of hair cells. The hairs project into a gelatinous cap called the cupula and monitor angular acceleration and deceleration from rotation. They would be stimulated by driving your car around a corner, turning your head, or falling forward. One canal lies horizontally, while the other two lie at about 45 degree angles to the horizontal axis. When the brain processes input from all three canals together, it can detect angular acceleration or deceleration in three dimensions. When the head turns, the fluid in the canals shifts, thereby bending stereocilia and sending signals to the brain. Upon cessation accelerating or decelerating—or just moving—the movement of the fluid within the canals slows or stops. For example, imagine holding a glass of water. When moving forward, water may splash backwards onto the hand, and when motion has stopped, water may splash forward onto the fingers. While in motion, the water settles in the glass and does not splash. Note that the canals are not sensitive to velocity itself, but to changes in velocity, so moving forward at 60mph with your eyes closed would not give the sensation of movement, but suddenly accelerating or braking would stimulate the receptors.
Higher Processing
Hair cells from the utricle, saccule, and semicircular canals also communicate through bipolar neurons to the cochlear nucleus in the medulla. Cochlear neurons send descending projections to the spinal cord and ascending projections to the pons, thalamus, and cerebellum. Connections to the cerebellum are important for coordinated movements. There are also projections to the temporal cortex, which account for feelings of dizziness; projections to autonomic nervous system areas in the brainstem, which account for motion sickness; and projections to the primary somatosensory cortex, which monitors subjective measurements of the external world and self-movement. People with lesions in the vestibular area of the somatosensory cortex see vertical objects in the world as being tilted. Finally, the vestibular signals project to certain optic muscles to coordinate eye and head movements.
Link to Learning
Click through this interactive tutorial to review the parts of the ear and how they function to process sound.
Summary
Audition is important for territory defense, predation, predator defense, and communal exchanges. The vestibular system, which is not auditory, detects linear acceleration and angular acceleration and deceleration. Both the auditory system and vestibular system use hair cells as their receptors.
Auditory stimuli are sound waves. The sound wave energy reaches the outer ear (pinna, canal, tympanum), and vibrations of the tympanum send the energy to the middle ear. The middle ear bones shift and the stapes transfers mechanical energy to the oval window of the fluid-filled inner ear cochlea. Once in the cochlea, the energy causes the basilar membrane to flex, thereby bending the stereocilia on receptor hair cells. This activates the receptors, which send their auditory neural signals to the brain.
The vestibular system has five parts that work together to provide the sense of direction, thus helping to maintain balance. The utricle and saccule measure head orientation: their calcium carbonate crystals shift when the head is tilted, thereby activating hair cells. The semicircular canals work similarly, such that when the head is turned, the fluid in the canals bends stereocilia on hair cells. The vestibular hair cells also send signals to the thalamus and to somatosensory cortex, but also to the cerebellum, the structure above the brainstem that plays a large role in timing and coordination of movement.
Glossary
audition
sense of hearing
basilar membrane
stiff structure in the cochlea that indirectly anchors auditory receptors
cochlea
whorled structure that contains receptors for transduction of the mechanical wave into an electrical signal
incus
(also, anvil) second of the three bones of the middle ear
inner ear
innermost part of the ear; consists of the cochlea and the vestibular system
labyrinth
bony, hollow structure that is the most internal part of the ear; contains the sites of transduction of auditory and vestibular information
malleus
(also, hammer) first of the three bones of the middle ear
middle ear
part of the hearing apparatus that functions to transfer energy from the tympanum to the oval window of the inner ear
organ of Corti
in the basilar membrane, the site of the transduction of sound, a mechanical wave, to a neural signal
ossicle
one of the three bones of the middle ear
outer ear
part of the ear that consists of the pinna, ear canal, and tympanum and which conducts sound waves into the middle ear
oval window
thin diaphragm between the middle and inner ears that receives sound waves from contact with the stapes bone of the middle ear
pinna
cartilaginous outer ear
semicircular canal
one of three half-circular, fluid-filled tubes in the vestibular labyrinth that monitors angular acceleration and deceleration
stapes
(also, stirrup) third of the three bones of the middle ear
stereocilia
in the auditory system, hair-like projections from hair cells that help detect sound waves
tectorial membrane
cochlear structure that lies above the hair cells and participates in the transduction of sound at the hair cells
tympanum
(also, tympanic membrane or ear drum) thin diaphragm between the outer and middle ears
ultrasound
sound frequencies above the human detectable ceiling of approximately 20,000 Hz | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/43%3A_Sensory_Systems/43.04%3A_Mechanoreceptors_2-_Hearing_Vibration_and_Balance/43.4.01%3A_Hearing_and_Vestibular_Sensation.txt |
Skills to Develop
• Explain in what way smell and taste stimuli differ from other sensory stimuli
• Identify the five primary tastes that can be distinguished by humans
• Explain in anatomical terms why a dog’s sense of smell is more acute than a human’s
Taste, also called gustation, and smell, also called olfaction, are the most interconnected senses in that both involve molecules of the stimulus entering the body and bonding to receptors. Smell lets an animal sense the presence of food or other animals—whether potential mates, predators, or prey—or other chemicals in the environment that can impact their survival. Similarly, the sense of taste allows animals to discriminate between types of foods. While the value of a sense of smell is obvious, what is the value of a sense of taste? Different tasting foods have different attributes, both helpful and harmful. For example, sweet-tasting substances tend to be highly caloric, which could be necessary for survival in lean times. Bitterness is associated with toxicity, and sourness is associated with spoiled food. Salty foods are valuable in maintaining homeostasis by helping the body retain water and by providing ions necessary for cells to function.
Tastes and Odors
Both taste and odor stimuli are molecules taken in from the environment. The primary tastes detected by humans are sweet, sour, bitter, salty and umami. The first four tastes need little explanation. The identification of umami as a fundamental taste occurred fairly recently—it was identified in 1908 by Japanese scientist Kikunae Ikeda while he worked with seaweed broth, but it was not widely accepted as a taste that could be physiologically distinguished until many years later. The taste of umami, also known as savoriness, is attributable to the taste of the amino acid L-glutamate. In fact, monosodium glutamate, or MSG, is often used in cooking to enhance the savory taste of certain foods. What is the adaptive value of being able to distinguish umami? Savory substances tend to be high in protein.
All odors that we perceive are molecules in the air we breathe. If a substance does not release molecules into the air from its surface, it has no smell. And if a human or other animal does not have a receptor that recognizes a specific molecule, then that molecule has no smell. Humans have about 350 olfactory receptor subtypes that work in various combinations to allow us to sense about 10,000 different odors. Compare that to mice, for example, which have about 1,300 olfactory receptor types, and therefore probably sense more odors. Both odors and tastes involve molecules that stimulate specific chemoreceptors. Although humans commonly distinguish taste as one sense and smell as another, they work together to create the perception of flavor. A person’s perception of flavor is reduced if he or she has congested nasal passages.
Reception and Transduction
Odorants (odor molecules) enter the nose and dissolve in the olfactory epithelium, the mucosa at the back of the nasal cavity (as illustrated in Figure \(1\)). The olfactory epithelium is a collection of specialized olfactory receptors in the back of the nasal cavity that spans an area about 5 cm2 in humans. Recall that sensory cells are neurons. An olfactory receptor, which is a dendrite of a specialized neuron, responds when it binds certain molecules inhaled from the environment by sending impulses directly to the olfactory bulb of the brain. Humans have about 12 million olfactory receptors, distributed among hundreds of different receptor types that respond to different odors. Twelve million seems like a large number of receptors, but compare that to other animals: rabbits have about 100 million, most dogs have about 1 billion, and bloodhounds—dogs selectively bred for their sense of smell—have about 4 billion. The overall size of the olfactory epithelium also differs between species, with that of bloodhounds, for example, being many times larger than that of humans.
Olfactory neurons are bipolar neurons (neurons with two processes from the cell body). Each neuron has a single dendrite buried in the olfactory epithelium, and extending from this dendrite are 5 to 20 receptor-laden, hair-like cilia that trap odorant molecules. The sensory receptors on the cilia are proteins, and it is the variations in their amino acid chains that make the receptors sensitive to different odorants. Each olfactory sensory neuron has only one type of receptor on its cilia, and the receptors are specialized to detect specific odorants, so the bipolar neurons themselves are specialized. When an odorant binds with a receptor that recognizes it, the sensory neuron associated with the receptor is stimulated. Olfactory stimulation is the only sensory information that directly reaches the cerebral cortex, whereas other sensations are relayed through the thalamus.
Evolution Connection: Pheromones
A pheromone is a chemical released by an animal that affects the behavior or physiology of animals of the same species. Pheromonal signals can have profound effects on animals that inhale them, but pheromones apparently are not consciously perceived in the same way as other odors. There are several different types of pheromones, which are released in urine or as glandular secretions. Certain pheromones are attractants to potential mates, others are repellants to potential competitors of the same sex, and still others play roles in mother-infant attachment. Some pheromones can also influence the timing of puberty, modify reproductive cycles, and even prevent embryonic implantation. While the roles of pheromones in many nonhuman species are important, pheromones have become less important in human behavior over evolutionary time compared to their importance to organisms with more limited behavioral repertoires.
The vomeronasal organ (VNO, or Jacobson’s organ) is a tubular, fluid-filled, olfactory organ present in many vertebrate animals that sits adjacent to the nasal cavity. It is very sensitive to pheromones and is connected to the nasal cavity by a duct. When molecules dissolve in the mucosa of the nasal cavity, they then enter the VNO where the pheromone molecules among them bind with specialized pheromone receptors. Upon exposure to pheromones from their own species or others, many animals, including cats, may display the flehmen response (Figure \(2\)), a curling of the upper lip that helps pheromone molecules enter the VNO.
Pheromonal signals are sent, not to the main olfactory bulb, but to a different neural structure that projects directly to the amygdala (recall that the amygdala is a brain center important in emotional reactions, such as fear). The pheromonal signal then continues to areas of the hypothalamus that are key to reproductive physiology and behavior. While some scientists assert that the VNO is apparently functionally vestigial in humans, even though there is a similar structure located near human nasal cavities, others are researching it as a possible functional system that may, for example, contribute to synchronization of menstrual cycles in women living in close proximity.
Taste
Detecting a taste (gustation) is fairly similar to detecting an odor (olfaction), given that both taste and smell rely on chemical receptors being stimulated by certain molecules. The primary organ of taste is the taste bud. A taste bud is a cluster of gustatory receptors (taste cells) that are located within the bumps on the tongue called papillae (singular: papilla) (illustrated in Figure \(3\)). There are several structurally distinct papillae. Filiform papillae, which are located across the tongue, are tactile, providing friction that helps the tongue move substances, and contain no taste cells. In contrast, fungiform papillae, which are located mainly on the anterior two-thirds of the tongue, each contain one to eight taste buds and also have receptors for pressure and temperature. The large circumvallate papillae contain up to 100 taste buds and form a V near the posterior margin of the tongue.
In addition to those two types of chemically and mechanically sensitive papillae are foliate papillae—leaf-like papillae located in parallel folds along the edges and toward the back of the tongue, as seen in the micrograph. Foliate papillae contain about 1,300 taste buds within their folds. Finally, there are circumvallate papillae, which are wall-like papillae in the shape of an inverted “V” at the back of the tongue. Each of these papillae is surrounded by a groove and contains about 250 taste buds.
Each taste bud’s taste cells are replaced every 10 to 14 days. These are elongated cells with hair-like processes called microvilli at the tips that extend into the taste bud pore (illustrated in Figure \(4\)). Food molecules (tastants) are dissolved in saliva, and they bind with and stimulate the receptors on the microvilli. The receptors for tastants are located across the outer portion and front of the tongue, outside of the middle area where the filiform papillae are most prominent.
In humans, there are five primary tastes, and each taste has only one corresponding type of receptor. Thus, like olfaction, each receptor is specific to its stimulus (tastant). Transduction of the five tastes happens through different mechanisms that reflect the molecular composition of the tastant. A salty tastant (containing NaCl) provides the sodium ions (Na+) that enter the taste neurons and excite them directly. Sour tastants are acids and belong to the thermoreceptor protein family. Binding of an acid or other sour-tasting molecule triggers a change in the ion channel and these increase hydrogen ion (H+) concentrations in the taste neurons, thus depolarizing them. Sweet, bitter, and umami tastants require a G-protein coupled receptor. These tastants bind to their respective receptors, thereby exciting the specialized neurons associated with them.
Both tasting abilities and sense of smell change with age. In humans, the senses decline dramatically by age 50 and continue to decline. A child may find a food to be too spicy, whereas an elderly person may find the same food to be bland and unappetizing.
Link to Learning
View this animation that shows how the sense of taste works.
Smell and Taste in the Brain
Olfactory neurons project from the olfactory epithelium to the olfactory bulb as thin, unmyelinated axons. The olfactory bulb is composed of neural clusters called glomeruli, and each glomerulus receives signals from one type of olfactory receptor, so each glomerulus is specific to one odorant. From glomeruli, olfactory signals travel directly to the olfactory cortex and then to the frontal cortex and the thalamus. Recall that this is a different path from most other sensory information, which is sent directly to the thalamus before ending up in the cortex. Olfactory signals also travel directly to the amygdala, thereafter reaching the hypothalamus, thalamus, and frontal cortex. The last structure that olfactory signals directly travel to is a cortical center in the temporal lobe structure important in spatial, autobiographical, declarative, and episodic memories. Olfaction is finally processed by areas of the brain that deal with memory, emotions, reproduction, and thought.
Taste neurons project from taste cells in the tongue, esophagus, and palate to the medulla, in the brainstem. From the medulla, taste signals travel to the thalamus and then to the primary gustatory cortex. Information from different regions of the tongue is segregated in the medulla, thalamus, and cortex.
Summary
There are five primary tastes in humans: sweet, sour, bitter, salty, and umami. Each taste has its own receptor type that responds only to that taste. Tastants enter the body and are dissolved in saliva. Taste cells are located within taste buds, which are found on three of the four types of papillae in the mouth.
Regarding olfaction, there are many thousands of odorants, but humans detect only about 10,000. Like taste receptors, olfactory receptors are each responsive to only one odorant. Odorants dissolve in nasal mucosa, where they excite their corresponding olfactory sensory cells. When these cells detect an odorant, they send their signals to the main olfactory bulb and then to other locations in the brain, including the olfactory cortex.
Glossary
bipolar neuron
neuron with two processes from the cell body, typically in opposite directions
glomerulus
in the olfactory bulb, one of the two neural clusters that receives signals from one type of olfactory receptor
gustation
sense of taste
odorant
airborne molecule that stimulates an olfactory receptor
olfaction
sense of smell
olfactory bulb
neural structure in the vertebrate brain that receives signals from olfactory receptors
olfactory epithelium
specialized tissue in the nasal cavity where olfactory receptors are located
olfactory receptor
dendrite of a specialized neuron
papilla
one of the small bump-like projections from the tongue
pheromone
substance released by an animal that can affect the physiology or behavior of other animals
tastant
food molecule that stimulates gustatory receptors
taste bud
clusters of taste cells
umami
one of the five basic tastes, which is described as “savory” and which may be largely the taste of L-glutamate | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/43%3A_Sensory_Systems/43.05%3A_Chemoreceptors-_Taste_Smell_and_pH.txt |
Skills to Develop
• Explain how electromagnetic waves differs from sound waves
• Trace the path of light through the eye to the point of the optic nerve
• Explain tonic activity as it is manifested in photoreceptors in the retina
Vision is the ability to detect light patterns from the outside environment and interpret them into images. Animals are bombarded with sensory information, and the sheer volume of visual information can be problematic. Fortunately, the visual systems of species have evolved to attend to the most-important stimuli. The importance of vision to humans is further substantiated by the fact that about one-third of the human cerebral cortex is dedicated to analyzing and perceiving visual information.
Light
As with auditory stimuli, light travels in waves. The compression waves that compose sound must travel in a medium—a gas, a liquid, or a solid. In contrast, light is composed of electromagnetic waves and needs no medium; light can travel in a vacuum (Figure \(1\)). The behavior of light can be discussed in terms of the behavior of waves and also in terms of the behavior of the fundamental unit of light—a packet of electromagnetic radiation called a photon. A glance at the electromagnetic spectrum shows that visible light for humans is just a small slice of the entire spectrum, which includes radiation that we cannot see as light because it is below the frequency of visible red light and above the frequency of visible violet light.
Certain variables are important when discussing perception of light. Wavelength (which varies inversely with frequency) manifests itself as hue. Light at the red end of the visible spectrum has longer wavelengths (and is lower frequency), while light at the violet end has shorter wavelengths (and is higher frequency). The wavelength of light is expressed in nanometers (nm); one nanometer is one billionth of a meter. Humans perceive light that ranges between approximately 380 nm and 740 nm. Some other animals, though, can detect wavelengths outside of the human range. For example, bees see near-ultraviolet light in order to locate nectar guides on flowers, and some non-avian reptiles sense infrared light (heat that prey gives off).
Wave amplitude is perceived as luminous intensity, or brightness. The standard unit of intensity of light is the candela, which is approximately the luminous intensity of a one common candle.
Light waves travel 299,792 km per second in a vacuum, (and somewhat slower in various media such as air and water), and those waves arrive at the eye as long (red), medium (green), and short (blue) waves. What is termed “white light” is light that is perceived as white by the human eye. This effect is produced by light that stimulates equally the color receptors in the human eye. The apparent color of an object is the color (or colors) that the object reflects. Thus a red object reflects the red wavelengths in mixed (white) light and absorbs all other wavelengths of light.
Anatomy of the Eye
The photoreceptive cells of the eye, where transduction of light to nervous impulses occurs, are located in the retina (shown in Figure \(2\)) on the inner surface of the back of the eye. But light does not impinge on the retina unaltered. It passes through other layers that process it so that it can be interpreted by the retina (Figure \(2\)b). The cornea, the front transparent layer of the eye, and the crystalline lens, a transparent convex structure behind the cornea, both refract (bend) light to focus the image on the retina. The iris, which is conspicuous as the colored part of the eye, is a circular muscular ring lying between the lens and cornea that regulates the amount of light entering the eye. In conditions of high ambient light, the iris contracts, reducing the size of the pupil at its center. In conditions of low light, the iris relaxes and the pupil enlarges.
Exercise
Which of the following statements about the human eye is false?
1. Rods detect color, while cones detect only shades of gray.
2. When light enters the retina, it passes the ganglion cells and bipolar cells before reaching photoreceptors at the rear of the eye.
3. The iris adjusts the amount of light coming into the eye.
4. The cornea is a protective layer on the front of the eye.
Answer
A
The main function of the lens is to focus light on the retina and fovea centralis. The lens is dynamic, focusing and re-focusing light as the eye rests on near and far objects in the visual field. The lens is operated by muscles that stretch it flat or allow it to thicken, changing the focal length of light coming through it to focus it sharply on the retina. With age comes the loss of the flexibility of the lens, and a form of farsightedness called presbyopia results. Presbyopia occurs because the image focuses behind the retina. Presbyopia is a deficit similar to a different type of farsightedness called hyperopia caused by an eyeball that is too short. For both defects, images in the distance are clear but images nearby are blurry. Myopia (nearsightedness) occurs when an eyeball is elongated and the image focus falls in front of the retina. In this case, images in the distance are blurry but images nearby are clear.
There are two types of photoreceptors in the retina: rods and cones, named for their general appearance as illustrated in Figure \(3\). Rods are strongly photosensitive and are located in the outer edges of the retina. They detect dim light and are used primarily for peripheral and nighttime vision. Cones are weakly photosensitive and are located near the center of the retina. They respond to bright light, and their primary role is in daytime, color vision.
The fovea is the region in the center back of the eye that is responsible for acute vision. The fovea has a high density of cones. When you bring your gaze to an object to examine it intently in bright light, the eyes orient so that the object’s image falls on the fovea. However, when looking at a star in the night sky or other object in dim light, the object can be better viewed by the peripheral vision because it is the rods at the edges of the retina, rather than the cones at the center, that operate better in low light. In humans, cones far outnumber rods in the fovea.
Link to Learning
Review the anatomical structure of the eye, clicking on each part to practice identification.
Transduction of Light
The rods and cones are the site of transduction of light to a neural signal. Both rods and cones contain photopigments. In vertebrates, the main photopigment, rhodopsin, has two main parts Figure \(4\)): an opsin, which is a membrane protein (in the form of a cluster of α-helices that span the membrane), and retinal—a molecule that absorbs light.
When light hits a photoreceptor, it causes a shape change in the retinal, altering its structure from a bent (cis) form of the molecule to its linear (trans) isomer. This isomerization of retinal activates the rhodopsin, starting a cascade of events that ends with the closing of Na+ channels in the membrane of the photoreceptor. Thus, unlike most other sensory neurons (which become depolarized by exposure to a stimulus) visual receptors become hyperpolarized and thus driven away from threshold (Figure \(5\)).
Trichromatic Coding
There are three types of cones (with different photopsins), and they differ in the wavelength to which they are most responsive, as shown in Figure \(6\). Some cones are maximally responsive to short light waves of 420 nm, so they are called S cones (“S” for “short”); others respond maximally to waves of 530 nm (M cones, for “medium”); a third group responds maximally to light of longer wavelengths, at 560 nm (L, or “long” cones). With only one type of cone, color vision would not be possible, and a two-cone (dichromatic) system has limitations. Primates use a three-cone (trichromatic) system, resulting in full color vision.
The color we perceive is a result of the ratio of activity of our three types of cones. The colors of the visual spectrum, running from long-wavelength light to short, are red (700 nm), orange (600 nm), yellow (565 nm), green (497 nm), blue (470 nm), indigo (450 nm), and violet (425 nm). Humans have very sensitive perception of color and can distinguish about 500 levels of brightness, 200 different hues, and 20 steps of saturation, or about 2 million distinct colors.
Retinal Processing
Visual signals leave the cones and rods, travel to the bipolar cells, and then to ganglion cells. A large degree of processing of visual information occurs in the retina itself, before visual information is sent to the brain.
Photoreceptors in the retina continuously undergo tonic activity. That is, they are always slightly active even when not stimulated by light. In neurons that exhibit tonic activity, the absence of stimuli maintains a firing rate at a baseline; while some stimuli increase firing rate from the baseline, and other stimuli decrease firing rate. In the absence of light, the bipolar neurons that connect rods and cones to ganglion cells are continuously and actively inhibited by the rods and cones. Exposure of the retina to light hyperpolarizes the rods and cones and removes their inhibition of bipolar cells. The now active bipolar cells in turn stimulate the ganglion cells, which send action potentials along their axons (which leave the eye as the optic nerve). Thus, the visual system relies on change in retinal activity, rather than the absence or presence of activity, to encode visual signals for the brain. Sometimes horizontal cells carry signals from one rod or cone to other photoreceptors and to several bipolar cells. When a rod or cone stimulates a horizontal cell, the horizontal cell inhibits more distant photoreceptors and bipolar cells, creating lateral inhibition. This inhibition sharpens edges and enhances contrast in the images by making regions receiving light appear lighter and dark surroundings appear darker. Amacrine cells can distribute information from one bipolar cell to many ganglion cells.
You can demonstrate this using an easy demonstration to “trick” your retina and brain about the colors you are observing in your visual field. Look fixedly at Figure \(7\) for about 45 seconds. Then quickly shift your gaze to a sheet of blank white paper or a white wall. You should see an afterimage of the Norwegian flag in its correct colors. At this point, close your eyes for a moment, then reopen them, looking again at the white paper or wall; the afterimage of the flag should continue to appear as red, white, and blue. What causes this? According to an explanation called opponent process theory, as you gazed fixedly at the green, black, and yellow flag, your retinal ganglion cells that respond positively to green, black, and yellow increased their firing dramatically. When you shifted your gaze to the neutral white ground, these ganglion cells abruptly decreased their activity and the brain interpreted this abrupt downshift as if the ganglion cells were responding now to their “opponent” colors: red, white, and blue, respectively, in the visual field. Once the ganglion cells return to their baseline activity state, the false perception of color will disappear.
Higher Processing
The myelinated axons of ganglion cells make up the optic nerves. Within the nerves, different axons carry different qualities of the visual signal. Some axons constitute the magnocellular (big cell) pathway, which carries information about form, movement, depth, and differences in brightness. Other axons constitute the parvocellular (small cell) pathway, which carries information on color and fine detail. Some visual information projects directly back into the brain, while other information crosses to the opposite side of the brain. This crossing of optical pathways produces the distinctive optic chiasma (Greek, for “crossing”) found at the base of the brain and allows us to coordinate information from both eyes.
Once in the brain, visual information is processed in several places, and its routes reflect the complexity and importance of visual information to humans and other animals. One route takes the signals to the thalamus, which serves as the routing station for all incoming sensory impulses except olfaction. In the thalamus, the magnocellular and parvocellular distinctions remain intact, and there are different layers of the thalamus dedicated to each. When visual signals leave the thalamus, they travel to the primary visual cortex at the rear of the brain. From the visual cortex, the visual signals travel in two directions. One stream that projects to the parietal lobe, in the side of the brain, carries magnocellular (“where”) information. A second stream projects to the temporal lobe and carries both magnocellular (“where”) and parvocellular (“what”) information.
Another important visual route is a pathway from the retina to the superior colliculus in the midbrain, where eye movements are coordinated and integrated with auditory information. Finally, there is the pathway from the retina to the suprachiasmatic nucleus (SCN) of the hypothalamus. The SCN is a cluster of cells that is considered to be the body’s internal clock, which controls our circadian (day-long) cycle. The SCN sends information to the pineal gland, which is important in sleep/wake patterns and annual cycles.
Link to Learning
View this interactive presentation to review what you have learned about how vision functions.
Summary
Vision is the only photo responsive sense. Visible light travels in waves and is a very small slice of the electromagnetic radiation spectrum. Light waves differ based on their frequency (wavelength = hue) and amplitude (intensity = brightness).
In the vertebrate retina, there are two types of light receptors (photoreceptors): cones and rods. Cones, which are the source of color vision, exist in three forms—L, M, and S—and they are differentially sensitive to different wavelengths. Cones are located in the retina, along with the dim-light, achromatic receptors (rods). Cones are found in the fovea, the central region of the retina, whereas rods are found in the peripheral regions of the retina.
Visual signals travel from the eye over the axons of retinal ganglion cells, which make up the optic nerves. Ganglion cells come in several versions. Some ganglion cell axons carry information on form, movement, depth, and brightness, while other axons carry information on color and fine detail. Visual information is sent to the superior colliculi in the midbrain, where coordination of eye movements and integration of auditory information takes place. Visual information is also sent to the suprachiasmatic nucleus (SCN) of the hypothalamus, which plays a role in the circadian cycle.
Glossary
candela
(cd) unit of measurement of luminous intensity (brightness)
circadian
describes a time cycle about one day in length
cone
weakly photosensitive, chromatic, cone-shaped neuron in the fovea of the retina that detects bright light and is used in daytime color vision
cornea
transparent layer over the front of the eye that helps focus light waves
fovea
region in the center of the retina with a high density of photoreceptors and which is responsible for acute vision
hyperopia
(also, farsightedness) visual defect in which the image focus falls behind the retina, thereby making images in the distance clear, but close-up images blurry
iris
pigmented, circular muscle at the front of the eye that regulates the amount of light entering the eye
lens
transparent, convex structure behind the cornea that helps focus light waves on the retina
myopia
(also, nearsightedness) visual defect in which the image focus falls in front of the retina, thereby making images in the distance blurry, but close-up images clear
presbyopia
visual defect in which the image focus falls behind the retina, thereby making images in the distance clear, but close-up images blurry; caused by age-based changes in the lens
pupil
small opening though which light enters
retina
layer of photoreceptive and supporting cells on the inner surface of the back of the eye
rhodopsin
main photopigment in vertebrates
rod
strongly photosensitive, achromatic, cylindrical neuron in the outer edges of the retina that detects dim light and is used in peripheral and nighttime vision
superior colliculus
paired structure in the top of the midbrain, which manages eye movements and auditory integration
suprachiasmatic nucleus
cluster of cells in the hypothalamus that plays a role in the circadian cycle
tonic activity
in a neuron, slight continuous activity while at rest
vision
sense of sight | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/43%3A_Sensory_Systems/43.06%3A_Vision.txt |
• 44.1: Regulation of Body Processes by Chemical Messengers
Chemical signals are released by signaling cells in the form of small, usually volatile or soluble molecules called ligands. A ligand is a molecule that binds another specific molecule, in some cases, delivering a signal in the process. Ligands can thus be thought of as signaling molecules. Ligands interact with proteins in target cells, which are cells that are affected by chemical signals; these proteins are also called receptors.
• 44.2: Overview of Hormone Action
Hormones cause cellular changes by binding to receptors on target cells. The number of receptors on a target cell can increase or decrease in response to hormone activity. Hormones can affect cells directly through intracellular hormone receptors or indirectly through plasma membrane hormone receptors. Lipid-derived (soluble) hormones can enter the cell by diffusing across the plasma membrane and binding to DNA to regulate gene transcription.
• 44.3: The Pituitary and Hypothalamus- The Body's Control Centers
There are three basic types of hormones: lipid-derived, amino acid-derived, and peptide. Lipid-derived hormones are structurally similar to cholesterol and include steroid hormones such as estradiol and testosterone. Amino acid-derived hormones are relatively small molecules and include the adrenal hormones epinephrine and norepinephrine. Peptide hormones are polypeptide chains or proteins and include the pituitary hormones, antidiuretic hormone (vasopressin), and oxytocin.
• 44.4: The Major Peripheral Endocrine Glands
Hormones have a wide range of effects and modulate many different body processes. The key regulatory processes that will be examined here are those affecting the excretory system, the reproductive system, metabolism, blood calcium concentrations, growth, and the stress response.
• 44.5: Other Hormones and Their Effects
Both the endocrine and nervous systems use chemical signals to communicate and regulate the body's physiology. The endocrine system releases hormones that act on target cells to regulate development, growth, energy metabolism, reproduction, and many behaviors. The nervous system releases neurotransmitters or neurohormones that regulate neurons, muscle cells, and endocrine cells.
44: The Endocrine System
Skills to Develop
• Describe four types of signaling found in multicellular organisms
• Compare internal receptors with cell-surface receptors
• Recognize the relationship between a ligand’s structure and its mechanism of action
There are two kinds of communication in the world of living cells. Communication between cells is called intercellular signaling, and communication within a cell is called intracellular signaling. An easy way to remember the distinction is by understanding the Latin origin of the prefixes: inter- means "between" (for example, intersecting lines are those that cross each other) and intra- means "inside" (like intravenous).
Chemical signals are released by signaling cells in the form of small, usually volatile or soluble molecules called ligands. A ligand is a molecule that binds another specific molecule, in some cases, delivering a signal in the process. Ligands can thus be thought of as signaling molecules. Ligands interact with proteins in target cells, which are cells that are affected by chemical signals; these proteins are also called receptors. Ligands and receptors exist in several varieties; however, a specific ligand will have a specific receptor that typically binds only that ligand.
Forms of Signaling
There are four categories of chemical signaling found in multicellular organisms: paracrine signaling, endocrine signaling, autocrine signaling, and direct signaling across gap junctions (Figure \(1\)). The main difference between the different categories of signaling is the distance that the signal travels through the organism to reach the target cell. Not all cells are affected by the same signals.
Paracrine Signaling
Signals that act locally between cells that are close together are called paracrine signals. Paracrine signals move by diffusion through the extracellular matrix. These types of signals usually elicit quick responses that last only a short amount of time. In order to keep the response localized, paracrine ligand molecules are normally quickly degraded by enzymes or removed by neighboring cells. Removing the signals will reestablish the concentration gradient for the signal, allowing them to quickly diffuse through the intracellular space if released again.
One example of paracrine signaling is the transfer of signals across synapses between nerve cells. A nerve cell consists of a cell body, several short, branched extensions called dendrites that receive stimuli, and a long extension called an axon, which transmits signals to other nerve cells or muscle cells. The junction between nerve cells where signal transmission occurs is called a synapse. A synaptic signal is a chemical signal that travels between nerve cells. Signals within the nerve cells are propagated by fast-moving electrical impulses. When these impulses reach the end of the axon, the signal continues on to a dendrite of the next cell by the release of chemical ligands called neurotransmitters by the presynaptic cell (the cell emitting the signal). The neurotransmitters are transported across the very small distances between nerve cells, which are called chemical synapses (Figure \(2\)). The small distance between nerve cells allows the signal to travel quickly; this enables an immediate response, such as, Take your hand off the stove!
When the neurotransmitter binds the receptor on the surface of the postsynaptic cell, the electrochemical potential of the target cell changes, and the next electrical impulse is launched. The neurotransmitters that are released into the chemical synapse are degraded quickly or get reabsorbed by the presynaptic cell so that the recipient nerve cell can recover quickly and be prepared to respond rapidly to the next synaptic signal.
Endocrine Signaling
Signals from distant cells are called endocrine signals, and they originate from endocrine cells. (In the body, many endocrine cells are located in endocrine glands, such as the thyroid gland, the hypothalamus, and the pituitary gland.) These types of signals usually produce a slower response but have a longer-lasting effect. The ligands released in endocrine signaling are called hormones, signaling molecules that are produced in one part of the body but affect other body regions some distance away.
Hormones travel the large distances between endocrine cells and their target cells via the bloodstream, which is a relatively slow way to move throughout the body. Because of their form of transport, hormones get diluted and are present in low concentrations when they act on their target cells. This is different from paracrine signaling, in which local concentrations of ligands can be very high.
Autocrine Signaling
Autocrine signals are produced by signaling cells that can also bind to the ligand that is released. This means the signaling cell and the target cell can be the same or a similar cell (the prefix auto- means self, a reminder that the signaling cell sends a signal to itself). This type of signaling often occurs during the early development of an organism to ensure that cells develop into the correct tissues and take on the proper function. Autocrine signaling also regulates pain sensation and inflammatory responses. Further, if a cell is infected with a virus, the cell can signal itself to undergo programmed cell death, killing the virus in the process. In some cases, neighboring cells of the same type are also influenced by the released ligand. In embryological development, this process of stimulating a group of neighboring cells may help to direct the differentiation of identical cells into the same cell type, thus ensuring the proper developmental outcome.
Direct Signaling Across Gap Junctions
Gap junctions in animals and plasmodesmata in plants are connections between the plasma membranes of neighboring cells. These water-filled channels allow small signaling molecules, called intracellular mediators, to diffuse between the two cells. Small molecules, such as calcium ions (Ca2+), are able to move between cells, but large molecules like proteins and DNA cannot fit through the channels. The specificity of the channels ensures that the cells remain independent but can quickly and easily transmit signals. The transfer of signaling molecules communicates the current state of the cell that is directly next to the target cell; this allows a group of cells to coordinate their response to a signal that only one of them may have received. In plants, plasmodesmata are ubiquitous, making the entire plant into a giant, communication network.
Types of Receptors
Receptors are protein molecules in the target cell or on its surface that bind ligand. There are two types of receptors, internal receptors and cell-surface receptors.
Internal receptors
Internal receptors, also known as intracellular or cytoplasmic receptors, are found in the cytoplasm of the cell and respond to hydrophobic ligand molecules that are able to travel across the plasma membrane. Once inside the cell, many of these molecules bind to proteins that act as regulators of mRNA synthesis (transcription) to mediate gene expression. Gene expression is the cellular process of transforming the information in a cell's DNA into a sequence of amino acids, which ultimately forms a protein. When the ligand binds to the internal receptor, a conformational change is triggered that exposes a DNA-binding site on the protein. The ligand-receptor complex moves into the nucleus, then binds to specific regulatory regions of the chromosomal DNA and promotes the initiation of transcription (Figure \(3\)). Transcription is the process of copying the information in a cells DNA into a special form of RNA called messenger RNA (mRNA); the cell uses information in the mRNA (which moves out into the cytoplasm and associates with ribosomes) to link specific amino acids in the correct order, producing a protein. Internal receptors can directly influence gene expression without having to pass the signal on to other receptors or messengers.
Cell-Surface Receptors
Cell-surface receptors, also known as transmembrane receptors, are cell surface, membrane-anchored (integral) proteins that bind to external ligand molecules. This type of receptor spans the plasma membrane and performs signal transduction, in which an extracellular signal is converted into an intercellular signal. Ligands that interact with cell-surface receptors do not have to enter the cell that they affect. Cell-surface receptors are also called cell-specific proteins or markers because they are specific to individual cell types.
Because cell-surface receptor proteins are fundamental to normal cell functioning, it should come as no surprise that a malfunction in any one of these proteins could have severe consequences. Errors in the protein structures of certain receptor molecules have been shown to play a role in hypertension (high blood pressure), asthma, heart disease, and cancer.
Each cell-surface receptor has three main components: an external ligand-binding domain, a hydrophobic membrane-spanning region, and an intracellular domain inside the cell. The ligand-binding domain is also called the extracellular domain. The size and extent of each of these domains vary widely, depending on the type of receptor.
Evolution Connection: How Viruses Recognize a Host
Unlike living cells, many viruses do not have a plasma membrane or any of the structures necessary to sustain life. Some viruses are simply composed of an inert protein shell containing DNA or RNA. To reproduce, viruses must invade a living cell, which serves as a host, and then take over the hosts cellular apparatus. But how does a virus recognize its host?
Viruses often bind to cell-surface receptors on the host cell. For example, the virus that causes human influenza (flu) binds specifically to receptors on membranes of cells of the respiratory system. Chemical differences in the cell-surface receptors among hosts mean that a virus that infects a specific species (for example, humans) cannot infect another species (for example, chickens).
However, viruses have very small amounts of DNA or RNA compared to humans, and, as a result, viral reproduction can occur rapidly. Viral reproduction invariably produces errors that can lead to changes in newly produced viruses; these changes mean that the viral proteins that interact with cell-surface receptors may evolve in such a way that they can bind to receptors in a new host. Such changes happen randomly and quite often in the reproductive cycle of a virus, but the changes only matter if a virus with new binding properties comes into contact with a suitable host. In the case of influenza, this situation can occur in settings where animals and people are in close contact, such as poultry and swine farms.1 Once a virus jumps to a new host, it can spread quickly. Scientists watch newly appearing viruses (called emerging viruses) closely in the hope that such monitoring can reduce the likelihood of global viral epidemics.
Cell-surface receptors are involved in most of the signaling in multicellular organisms. There are three general categories of cell-surface receptors: ion channel-linked receptors, G-protein-linked receptors, and enzyme-linked receptors.
Ion channel-linked receptors bind a ligand and open a channel through the membrane that allows specific ions to pass through. To form a channel, this type of cell-surface receptor has an extensive membrane-spanning region. In order to interact with the phospholipid fatty acid tails that form the center of the plasma membrane, many of the amino acids in the membrane-spanning region are hydrophobic in nature. Conversely, the amino acids that line the inside of the channel are hydrophilic to allow for the passage of water or ions. When a ligand binds to the extracellular region of the channel, there is a conformational change in the proteins structure that allows ions such as sodium, calcium, magnesium, and hydrogen to pass through (Figure \(4\)).
G-protein-linked receptors bind a ligand and activate a membrane protein called a G-protein. The activated G-protein then interacts with either an ion channel or an enzyme in the membrane (Figure \(5\)). All G-protein-linked receptors have seven transmembrane domains, but each receptor has its own specific extracellular domain and G-protein-binding site.
Cell signaling using G-protein-linked receptors occurs as a cyclic series of events. Before the ligand binds, the inactive G-protein can bind to a newly revealed site on the receptor specific for its binding. Once the G-protein binds to the receptor, the resultant shape change activates the G-protein, which releases GDP and picks up GTP. The subunits of the G-protein then split into the α subunit and the βγ subunit. One or both of these G-protein fragments may be able to activate other proteins as a result. After awhile, the GTP on the active α subunit of the G-protein is hydrolyzed to GDP and the βγ subunit is deactivated. The subunits reassociate to form the inactive G-protein and the cycle begins anew.
G-protein-linked receptors have been extensively studied and much has been learned about their roles in maintaining health. Bacteria that are pathogenic to humans can release poisons that interrupt specific G-protein-linked receptor function, leading to illnesses such as pertussis, botulism, and cholera. In cholera (Figure \(6\)), for example, the water-borne bacterium Vibrio cholerae produces a toxin, choleragen, that binds to cells lining the small intestine. The toxin then enters these intestinal cells, where it modifies a G-protein that controls the opening of a chloride channel and causes it to remain continuously active, resulting in large losses of fluids from the body and potentially fatal dehydration as a result.
Enzyme-linked receptors are cell-surface receptors with intracellular domains that are associated with an enzyme. In some cases, the intracellular domain of the receptor itself is an enzyme. Other enzyme-linked receptors have a small intracellular domain that interacts directly with an enzyme. The enzyme-linked receptors normally have large extracellular and intracellular domains, but the membrane-spanning region consists of a single alpha-helical region of the peptide strand. When a ligand binds to the extracellular domain, a signal is transferred through the membrane, activating the enzyme. Activation of the enzyme sets off a chain of events within the cell that eventually leads to a response. One example of this type of enzyme-linked receptor is the tyrosine kinase receptor (Figure \(7\)). A kinase is an enzyme that transfers phosphate groups from ATP to another protein. The tyrosine kinase receptor transfers phosphate groups to tyrosine molecules (tyrosine residues). First, signaling molecules bind to the extracellular domain of two nearby tyrosine kinase receptors. The two neighboring receptors then bond together, or dimerize. Phosphates are then added to tyrosine residues on the intracellular domain of the receptors (phosphorylation). The phosphorylated residues can then transmit the signal to the next messenger within the cytoplasm.
Art Connection
HER2 is a receptor tyrosine kinase. In 30 percent of human breast cancers, HER2 is permanently activated, resulting in unregulated cell division. Lapatinib, a drug used to treat breast cancer, inhibits HER2 receptor tyrosine kinase autophosphorylation (the process by which the receptor adds phosphates onto itself), thus reducing tumor growth by 50 percent. Besides autophosphorylation, which of the following steps would be inhibited by Lapatinib?
1. Signaling molecule binding, dimerization, and the downstream cellular response
2. Dimerization, and the downstream cellular response
3. The downstream cellular response
4. Phosphatase activity, dimerization, and the downsteam cellular response
Signaling Molecules
Produced by signaling cells and the subsequent binding to receptors in target cells, ligands act as chemical signals that travel to the target cells to coordinate responses. The types of molecules that serve as ligands are incredibly varied and range from small proteins to small ions like calcium (Ca2+).
Small Hydrophobic Ligands
Small hydrophobic ligands can directly diffuse through the plasma membrane and interact with internal receptors. Important members of this class of ligands are the steroid hormones. Steroids are lipids that have a hydrocarbon skeleton with four fused rings; different steroids have different functional groups attached to the carbon skeleton. Steroid hormones include the female sex hormone, estradiol, which is a type of estrogen; the male sex hormone, testosterone; and cholesterol, which is an important structural component of biological membranes and a precursor of steroid hormones (Figure \(8\)). Other hydrophobic hormones include thyroid hormones and vitamin D. In order to be soluble in blood, hydrophobic ligands must bind to carrier proteins while they are being transported through the bloodstream.
Water-Soluble Ligands
Water-soluble ligands are polar and therefore cannot pass through the plasma membrane unaided; sometimes, they are too large to pass through the membrane at all. Instead, most water-soluble ligands bind to the extracellular domain of cell-surface receptors. This group of ligands is quite diverse and includes small molecules, peptides, and proteins.
Other Ligands
Nitric oxide (NO) is a gas that also acts as a ligand. It is able to diffuse directly across the plasma membrane, and one of its roles is to interact with receptors in smooth muscle and induce relaxation of the tissue. NO has a very short half-life and therefore only functions over short distances. Nitroglycerin, a treatment for heart disease, acts by triggering the release of NO, which causes blood vessels to dilate (expand), thus restoring blood flow to the heart. NO has become better known recently because the pathway that it affects is targeted by prescription medications for erectile dysfunction, such as Viagra (erection involves dilated blood vessels).
Summary
Cells communicate by both inter- and intracellular signaling. Signaling cells secrete ligands that bind to target cells and initiate a chain of events within the target cell. The four categories of signaling in multicellular organisms are paracrine signaling, endocrine signaling, autocrine signaling, and direct signaling across gap junctions. Paracrine signaling takes place over short distances. Endocrine signals are carried long distances through the bloodstream by hormones, and autocrine signals are received by the same cell that sent the signal or other nearby cells of the same kind. Gap junctions allow small molecules, including signaling molecules, to flow between neighboring cells.
Internal receptors are found in the cell cytoplasm. Here, they bind ligand molecules that cross the plasma membrane; these receptor-ligand complexes move to the nucleus and interact directly with cellular DNA. Cell-surface receptors transmit a signal from outside the cell to the cytoplasm. Ion channel-linked receptors, when bound to their ligands, form a pore through the plasma membrane through which certain ions can pass. G-protein-linked receptors interact with a G-protein on the cytoplasmic side of the plasma membrane, promoting the exchange of bound GDP for GTP and interacting with other enzymes or ion channels to transmit a signal. Enzyme-linked receptors transmit a signal from outside the cell to an intracellular domain of a membrane-bound enzyme. Ligand binding causes activation of the enzyme. Small hydrophobic ligands (like steroids) are able to penetrate the plasma membrane and bind to internal receptors. Water-soluble hydrophilic ligands are unable to pass through the membrane; instead, they bind to cell-surface receptors, which transmit the signal to the inside of the cell.
Art Connections
Figure \(7\): HER2 is a receptor tyrosine kinase. In 30 percent of human breast cancers, HER2 is permanently activated, resulting in unregulated cell division. Lapatinib, a drug used to treat breast cancer, inhibits HER2 receptor tyrosine kinase autophosphorylation (the process by which the receptor adds phosphates onto itself), thus reducing tumor growth by 50 percent. Besides autophosphorylation, which of the following steps would be inhibited by Lapatinib?
1. Signaling molecule binding, dimerization, and the downstream cellular response.
2. Dimerization, and the downstream cellular response.
3. The downstream cellular response.
4. Phosphatase activity, dimerization, and the downsteam cellular response.
Answer
C. The downstream cellular response would be inhibited.
Footnotes
1. 1 A. B. Sigalov, The School of Nature. IV. Learning from Viruses, Self/Nonself 1, no. 4 (2010): 282-298. Y. Cao, X. Koh, L. Dong, X. Du, A. Wu, X. Ding, H. Deng, Y. Shu, J. Chen, T. Jiang, Rapid Estimation of Binding Activity of Influenza Virus Hemagglutinin to Human and Avian Receptors, PLoS One 6, no. 4 (2011): e18664.
Glossary
autocrine signal
signal that is sent and received by the same or similar nearby cells
cell-surface receptor
cell-surface protein that transmits a signal from the exterior of the cell to the interior, even though the ligand does not enter the cell
chemical synapse
small space between axon terminals and dendrites of nerve cells where neurotransmitters function
endocrine cell
cell that releases ligands involved in endocrine signaling (hormones)
endocrine signal
long-distance signal that is delivered by ligands (hormones) traveling through an organisms circulatory system from the signaling cell to the target cell
enzyme-linked receptor
cell-surface receptor with intracellular domains that are associated with membrane-bound enzymes
extracellular domain
region of a cell-surface receptor that is located on the cell surface
G-protein-linked receptor
cell-surface receptor that activates membrane-bound G-proteins to transmit a signal from the receptor to nearby membrane components
intercellular signaling
communication between cells
internal receptor
(also, intracellular receptor) receptor protein that is located in the cytosol of a cell and binds to ligands that pass through the plasma membrane
intracellular mediator
(also, second messenger) small molecule that transmits signals within a cell
intracellular signaling
communication within cells
ion channel-linked receptor
cell-surface receptor that forms a plasma membrane channel, which opens when a ligand binds to the extracellular domain (ligand-gated channels)
ligand
molecule produced by a signaling cell that binds with a specific receptor, delivering a signal in the process
neurotransmitter
chemical ligand that carries a signal from one nerve cell to the next
paracrine signal
signal between nearby cells that is delivered by ligands traveling in the liquid medium in the space between the cells
receptor
protein in or on a target cell that bind to ligands
signaling cell
cell that releases signal molecules that allow communication with another cell
synaptic signal
chemical signal (neurotransmitter) that travels between nerve cells
target cell
cell that has a receptor for a signal or ligand from a signaling cell
44.01: Regulation of Body Processes by Chemical Messengers
Skills to Develop
• List the different types of hormones
• Explain their role in maintaining homeostasis
Maintaining homeostasis within the body requires the coordination of many different systems and organs. Communication between neighboring cells, and between cells and tissues in distant parts of the body, occurs through the release of chemicals called hormones. Hormones are released into body fluids (usually blood) that carry these chemicals to their target cells. At the target cells, which are cells that have a receptor for a signal or ligand from a signal cell, the hormones elicit a response. The cells, tissues, and organs that secrete hormones make up the endocrine system. Examples of glands of the endocrine system include the adrenal glands, which produce hormones such as epinephrine and norepinephrine that regulate responses to stress, and the thyroid gland, which produces thyroid hormones that regulate metabolic rates.
Although there are many different hormones in the human body, they can be divided into three classes based on their chemical structure: lipid-derived, amino acid-derived, and peptide (peptide and proteins) hormones. One of the key distinguishing features of lipid-derived hormones is that they can diffuse across plasma membranes whereas the amino acid-derived and peptide hormones cannot.
Lipid-Derived Hormones (or Lipid-soluble Hormones)
Most lipid hormones are derived from cholesterol and thus are structurally similar to it, as illustrated in Figure \(1\). The primary class of lipid hormones in humans is the steroid hormones. Chemically, these hormones are usually ketones or alcohols; their chemical names will end in “-ol” for alcohols or “-one” for ketones. Examples of steroid hormones include estradiol, which is an estrogen, or female sex hormone, and testosterone, which is an androgen, or male sex hormone. These two hormones are released by the female and male reproductive organs, respectively. Other steroid hormones include aldosterone and cortisol, which are released by the adrenal glands along with some other types of androgens. Steroid hormones are insoluble in water, and they are transported by transport proteins in blood. As a result, they remain in circulation longer than peptide hormones. For example, cortisol has a half-life of 60 to 90 minutes, while epinephrine, an amino acid derived-hormone, has a half-life of approximately one minute.
Amino Acid-Derived Hormones
The amino acid-derived hormones are relatively small molecules that are derived from the amino acids tyrosine and tryptophan, shown in Figure \(2\). If a hormone is amino acid-derived, its chemical name will end in “-ine”. Examples of amino acid-derived hormones include epinephrine and norepinephrine, which are synthesized in the medulla of the adrenal glands, and thyroxine, which is produced by the thyroid gland. The pineal gland in the brain makes and secretes melatonin which regulates sleep cycles.
Peptide Hormones
The structure of peptide hormones is that of a polypeptide chain (chain of amino acids). The peptide hormones include molecules that are short polypeptide chains, such as antidiuretic hormone and oxytocin produced in the brain and released into the blood in the posterior pituitary gland. This class also includes small proteins, like growth hormones produced by the pituitary, and large glycoproteins such as follicle-stimulating hormone produced by the pituitary. Figure \(3\) illustrates these peptide hormones.
Secreted peptides like insulin are stored within vesicles in the cells that synthesize them. They are then released in response to stimuli such as high blood glucose levels in the case of insulin. Amino acid-derived and polypeptide hormones are water-soluble and insoluble in lipids. These hormones cannot pass through plasma membranes of cells; therefore, their receptors are found on the surface of the target cells.
Career Connection: Endocrinologist
An endocrinologist is a medical doctor who specializes in treating disorders of the endocrine glands, hormone systems, and glucose and lipid metabolic pathways. An endocrine surgeon specializes in the surgical treatment of endocrine diseases and glands. Some of the diseases that are managed by endocrinologists: disorders of the pancreas (diabetes mellitus), disorders of the pituitary (gigantism, acromegaly, and pituitary dwarfism), disorders of the thyroid gland (goiter and Graves’ disease), and disorders of the adrenal glands (Cushing’s disease and Addison’s disease).
Endocrinologists are required to assess patients and diagnose endocrine disorders through extensive use of laboratory tests. Many endocrine diseases are diagnosed using tests that stimulate or suppress endocrine organ functioning. Blood samples are then drawn to determine the effect of stimulating or suppressing an endocrine organ on the production of hormones. For example, to diagnose diabetes mellitus, patients are required to fast for 12 to 24 hours. They are then given a sugary drink, which stimulates the pancreas to produce insulin to decrease blood glucose levels. A blood sample is taken one to two hours after the sugar drink is consumed. If the pancreas is functioning properly, the blood glucose level will be within a normal range. Another example is the A1C test, which can be performed during blood screening. The A1C test measures average blood glucose levels over the past two to three months by examining how well the blood glucose is being managed over a long time.
Once a disease has been diagnosed, endocrinologists can prescribe lifestyle changes and/or medications to treat the disease. Some cases of diabetes mellitus can be managed by exercise, weight loss, and a healthy diet; in other cases, medications may be required to enhance insulin release. If the disease cannot be controlled by these means, the endocrinologist may prescribe insulin injections.
In addition to clinical practice, endocrinologists may also be involved in primary research and development activities. For example, ongoing islet transplant research is investigating how healthy pancreas islet cells may be transplanted into diabetic patients. Successful islet transplants may allow patients to stop taking insulin injections.
Summary
There are three basic types of hormones: lipid-derived, amino acid-derived, and peptide. Lipid-derived hormones are structurally similar to cholesterol and include steroid hormones such as estradiol and testosterone. Amino acid-derived hormones are relatively small molecules and include the adrenal hormones epinephrine and norepinephrine. Peptide hormones are polypeptide chains or proteins and include the pituitary hormones, antidiuretic hormone (vasopressin), and oxytocin.
Glossary
amino acid-derived hormone
hormone derived from amino acids
lipid-derived hormone
hormone derived mostly from cholesterol
peptide hormone
hormone composed of a polypeptide chain | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.01%3A_Regulation_of_Body_Processes_by_Chemical_Messengers/44.1.01%3A_Types_of_Hormones.txt |
Skills to Develop
• Explain how hormones regulate the excretory system
• Discuss the role of hormones in the reproductive system
• Describe how hormones regulate metabolism
• Explain the role of hormones in different diseases
Hormones have a wide range of effects and modulate many different body processes. The key regulatory processes that will be examined here are those affecting the excretory system, the reproductive system, metabolism, blood calcium concentrations, growth, and the stress response.
Hormonal Regulation of the Excretory System
Maintaining a proper water balance in the body is important to avoid dehydration or over-hydration (hyponatremia). The water concentration of the body is monitored by osmoreceptors in the hypothalamus, which detect the concentration of electrolytes in the extracellular fluid. The concentration of electrolytes in the blood rises when there is water loss caused by excessive perspiration, inadequate water intake, or low blood volume due to blood loss. An increase in blood electrolyte levels results in a neuronal signal being sent from the osmoreceptors in hypothalamic nuclei. The pituitary gland has two components: anterior and posterior. The anterior pituitary is composed of glandular cells that secrete protein hormones. The posterior pituitary is an extension of the hypothalamus. It is composed largely of neurons that are continuous with the hypothalamus.
The hypothalamus produces a polypeptide hormone known as antidiuretic hormone (ADH), which is transported to and released from the posterior pituitary gland. The principal action of ADH is to regulate the amount of water excreted by the kidneys. As ADH (which is also known as vasopressin) causes direct water reabsorption from the kidney tubules, salts and wastes are concentrated in what will eventually be excreted as urine. The hypothalamus controls the mechanisms of ADH secretion, either by regulating blood volume or the concentration of water in the blood. Dehydration or physiological stress can cause an increase of osmolarity above 300 mOsm/L, which in turn, raises ADH secretion and water will be retained, causing an increase in blood pressure. ADH travels in the bloodstream to the kidneys. Once at the kidneys, ADH changes the kidneys to become more permeable to water by temporarily inserting water channels, aquaporins, into the kidney tubules. Water moves out of the kidney tubules through the aquaporins, reducing urine volume. The water is reabsorbed into the capillaries lowering blood osmolarity back toward normal. As blood osmolarity decreases, a negative feedback mechanism reduces osmoreceptor activity in the hypothalamus, and ADH secretion is reduced. ADH release can be reduced by certain substances, including alcohol, which can cause increased urine production and dehydration.
Chronic underproduction of ADH or a mutation in the ADH receptor results in diabetes insipidus. If the posterior pituitary does not release enough ADH, water cannot be retained by the kidneys and is lost as urine. This causes increased thirst, but water taken in is lost again and must be continually consumed. If the condition is not severe, dehydration may not occur, but severe cases can lead to electrolyte imbalances due to dehydration.
Another hormone responsible for maintaining electrolyte concentrations in extracellular fluids is aldosterone, a steroid hormone that is produced by the adrenal cortex. In contrast to ADH, which promotes the reabsorption of water to maintain proper water balance, aldosterone maintains proper water balance by enhancing Na+ reabsorption and K+ secretion from extracellular fluid of the cells in kidney tubules. Because it is produced in the cortex of the adrenal gland and affects the concentrations of minerals Na+ and K+, aldosterone is referred to as a mineralocorticoid, a corticosteroid that affects ion and water balance. Aldosterone release is stimulated by a decrease in blood sodium levels, blood volume, or blood pressure, or an increase in blood potassium levels. It also prevents the loss of Na+ from sweat, saliva, and gastric juice. The reabsorption of Na+ also results in the osmotic reabsorption of water, which alters blood volume and blood pressure.
Aldosterone production can be stimulated by low blood pressure, which triggers a sequence of chemical release, as illustrated in Figure \(1\). When blood pressure drops, the renin-angiotensin-aldosterone system (RAAS) is activated. Cells in the juxtaglomerular apparatus, which regulates the functions of the nephrons of the kidney, detect this and release renin. Renin, an enzyme, circulates in the blood and reacts with a plasma protein produced by the liver called angiotensinogen. When angiotensinogen is cleaved by renin, it produces angiotensin I, which is then converted into angiotensin II in the lungs. Angiotensin II functions as a hormone and then causes the release of the hormone aldosterone by the adrenal cortex, resulting in increased Na+ reabsorption, water retention, and an increase in blood pressure. Angiotensin II in addition to being a potent vasoconstrictor also causes an increase in ADH and increased thirst, both of which help to raise blood pressure.
Hormonal Regulation of the Reproductive System
Regulation of the reproductive system is a process that requires the action of hormones from the pituitary gland, the adrenal cortex, and the gonads. During puberty in both males and females, the hypothalamus produces gonadotropin-releasing hormone (GnRH), which stimulates the production and release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary gland. These hormones regulate the gonads (testes in males and ovaries in females) and therefore are called gonadotropins. In both males and females, FSH stimulates gamete production and LH stimulates production of hormones by the gonads. An increase in gonad hormone levels inhibits GnRH production through a negative feedback loop.
Regulation of the Male Reproductive System
In males, FSH stimulates the maturation of sperm cells. FSH production is inhibited by the hormone inhibin, which is released by the testes. LH stimulates production of the sex hormones (androgens) by the interstitial cells of the testes and therefore is also called interstitial cell-stimulating hormone. The most widely known androgen in males is testosterone. Testosterone promotes the production of sperm and masculine characteristics. The adrenal cortex also produces small amounts of testosterone precursor, although the role of this additional hormone production is not fully understood.
Everyday Connection: The Dangers of Synthetic Hormones
Some athletes attempt to boost their performance by using artificial hormones that enhance muscle performance. Anabolic steroids, a form of the male sex hormone testosterone, are one of the most widely known performance-enhancing drugs. Steroids are used to help build muscle mass. Other hormones that are used to enhance athletic performance include erythropoietin, which triggers the production of red blood cells, and human growth hormone, which can help in building muscle mass. Most performance enhancing drugs are illegal for non-medical purposes. They are also banned by national and international governing bodies including the International Olympic Committee, the U.S. Olympic Committee, the National Collegiate Athletic Association, the Major League Baseball, and the National Football League.
The side effects of synthetic hormones are often significant and non-reversible, and in some cases, fatal. Androgens produce several complications such as liver dysfunctions and liver tumors, prostate gland enlargement, difficulty urinating, premature closure of epiphyseal cartilages, testicular atrophy, infertility, and immune system depression. The physiological strain caused by these substances is often greater than what the body can handle, leading to unpredictable and dangerous effects and linking their use to heart attacks, strokes, and impaired cardiac function.
Regulation of the Female Reproductive System
In females, FSH stimulates development of egg cells, called ova, which develop in structures called follicles. Follicle cells produce the hormone inhibin, which inhibits FSH production. LH also plays a role in the development of ova, induction of ovulation, and stimulation of estradiol and progesterone production by the ovaries, as illustrated in Figure \(3\). Estradiol and progesterone are steroid hormones that prepare the body for pregnancy. Estradiol produces secondary sex characteristics in females, while both estradiol and progesterone regulate the menstrual cycle.
In addition to producing FSH and LH, the anterior portion of the pituitary gland also produces the hormone prolactin (PRL) in females. Prolactin stimulates the production of milk by the mammary glands following childbirth. Prolactin levels are regulated by the hypothalamic hormones prolactin-releasing hormone (PRH) and prolactin-inhibiting hormone (PIH), which is now known to be dopamine. PRH stimulates the release of prolactin and PIH inhibits it.
The posterior pituitary releases the hormone oxytocin, which stimulates uterine contractions during childbirth. The uterine smooth muscles are not very sensitive to oxytocin until late in pregnancy when the number of oxytocin receptors in the uterus peaks. Stretching of tissues in the uterus and cervix stimulates oxytocin release during childbirth. Contractions increase in intensity as blood levels of oxytocin rise via a positive feedback mechanism until the birth is complete. Oxytocin also stimulates the contraction of myoepithelial cells around the milk-producing mammary glands. As these cells contract, milk is forced from the secretory alveoli into milk ducts and is ejected from the breasts in milk ejection (“let-down”) reflex. Oxytocin release is stimulated by the suckling of an infant, which triggers the synthesis of oxytocin in the hypothalamus and its release into circulation at the posterior pituitary.
Hormonal Regulation of Metabolism
Blood glucose levels vary widely over the course of a day as periods of food consumption alternate with periods of fasting. Insulin and glucagon are the two hormones primarily responsible for maintaining homeostasis of blood glucose levels. Additional regulation is mediated by the thyroid hormones.
Regulation of Blood Glucose Levels by Insulin and Glucagon
Cells of the body require nutrients in order to function, and these nutrients are obtained through feeding. In order to manage nutrient intake, storing excess intake and utilizing reserves when necessary, the body uses hormones to moderate energy stores. Insulin is produced by the beta cells of the pancreas, which are stimulated to release insulin as blood glucose levels rise (for example, after a meal is consumed). Insulin lowers blood glucose levels by enhancing the rate of glucose uptake and utilization by target cells, which use glucose for ATP production. It also stimulates the liver to convert glucose to glycogen, which is then stored by cells for later use. Insulin also increases glucose transport into certain cells, such as muscle cells and the liver. This results from an insulin-mediated increase in the number of glucose transporter proteins in cell membranes, which remove glucose from circulation by facilitated diffusion. As insulin binds to its target cell via insulin receptors and signal transduction, it triggers the cell to incorporate glucose transport proteins into its membrane. This allows glucose to enter the cell, where it can be used as an energy source. However, this does not occur in all cells: some cells, including those in the kidneys and brain, can access glucose without the use of insulin. Insulin also stimulates the conversion of glucose to fat in adipocytes and the synthesis of proteins. These actions mediated by insulin cause blood glucose concentrations to fall, called a hypoglycemic “low sugar” effect, which inhibits further insulin release from beta cells through a negative feedback loop.
Link to Learning
This animation describes the role of insulin and the pancreas in diabetes.
Impaired insulin function can lead to a condition called diabetes mellitus, the main symptoms of which are illustrated in Figure \(4\). This can be caused by low levels of insulin production by the beta cells of the pancreas, or by reduced sensitivity of tissue cells to insulin. This prevents glucose from being absorbed by cells, causing high levels of blood glucose, or hyperglycemia (high sugar). High blood glucose levels make it difficult for the kidneys to recover all the glucose from nascent urine, resulting in glucose being lost in urine. High glucose levels also result in less water being reabsorbed by the kidneys, causing high amounts of urine to be produced; this may result in dehydration. Over time, high blood glucose levels can cause nerve damage to the eyes and peripheral body tissues, as well as damage to the kidneys and cardiovascular system. Oversecretion of insulin can cause hypoglycemia, low blood glucose levels. This causes insufficient glucose availability to cells, often leading to muscle weakness, and can sometimes cause unconsciousness or death if left untreated.
When blood glucose levels decline below normal levels, for example between meals or when glucose is utilized rapidly during exercise, the hormone glucagon is released from the alpha cells of the pancreas. Glucagon raises blood glucose levels, eliciting what is called a hyperglycemic effect, by stimulating the breakdown of glycogen to glucose in skeletal muscle cells and liver cells in a process called glycogenolysis. Glucose can then be utilized as energy by muscle cells and released into circulation by the liver cells. Glucagon also stimulates absorption of amino acids from the blood by the liver, which then converts them to glucose. This process of glucose synthesis is called gluconeogenesis. Glucagon also stimulates adipose cells to release fatty acids into the blood. These actions mediated by glucagon result in an increase in blood glucose levels to normal homeostatic levels. Rising blood glucose levels inhibit further glucagon release by the pancreas via a negative feedback mechanism. In this way, insulin and glucagon work together to maintain homeostatic glucose levels, as shown in Figure \(5\).
Exercise
Pancreatic tumors may cause excess secretion of glucagon. Type I diabetes results from the failure of the pancreas to produce insulin. Which of the following statement about these two conditions is true?
1. A pancreatic tumor and type I diabetes will have the opposite effects on blood sugar levels.
2. A pancreatic tumor and type I diabetes will both cause hyperglycemia.
3. A pancreatic tumor and type I diabetes will both cause hypoglycemia.
4. Both pancreatic tumors and type I diabetes result in the inability of cells to take up glucose.
Answer
B
Regulation of Blood Glucose Levels by Thyroid Hormones
The basal metabolic rate, which is the amount of calories required by the body at rest, is determined by two hormones produced by the thyroid gland: thyroxine, also known as tetraiodothyronine or T4, and triiodothyronine, also known as T3. These hormones affect nearly every cell in the body except for the adult brain, uterus, testes, blood cells, and spleen. They are transported across the plasma membrane of target cells and bind to receptors on the mitochondria resulting in increased ATP production. In the nucleus, T3 and T4 activate genes involved in energy production and glucose oxidation. This results in increased rates of metabolism and body heat production, which is known as the hormone’s calorigenic effect.
T3 and T4 release from the thyroid gland is stimulated by thyroid-stimulating hormone (TSH), which is produced by the anterior pituitary. TSH binding at the receptors of the follicle of the thyroid triggers the production of T3 and T4 from a glycoprotein called thyroglobulin. Thyroglobulin is present in the follicles of the thyroid, and is converted into thyroid hormones with the addition of iodine. Iodine is formed from iodide ions that are actively transported into the thyroid follicle from the bloodstream. A peroxidase enzyme then attaches the iodine to the tyrosine amino acid found in thyroglobulin. T3 has three iodine ions attached, while T4 has four iodine ions attached. T3 and T4 are then released into the bloodstream, with T4 being released in much greater amounts than T3. As T3 is more active than T4 and is responsible for most of the effects of thyroid hormones, tissues of the body convert T4 to T3 by the removal of an iodine ion. Most of the released T3 and T4 becomes attached to transport proteins in the bloodstream and is unable to cross the plasma membrane of cells. These protein-bound molecules are only released when blood levels of the unattached hormone begin to decline. In this way, a week’s worth of reserve hormone is maintained in the blood. Increased T3 and T4 levels in the blood inhibit the release of TSH, which results in lower T3 and T4 release from the thyroid.
The follicular cells of the thyroid require iodides (anions of iodine) in order to synthesize T3 and T4. Iodides obtained from the diet are actively transported into follicle cells resulting in a concentration that is approximately 30 times higher than in blood. The typical diet in North America provides more iodine than required due to the addition of iodide to table salt. Inadequate iodine intake, which occurs in many developing countries, results in an inability to synthesize T3 and T4 hormones. The thyroid gland enlarges in a condition called goiter, which is caused by overproduction of TSH without the formation of thyroid hormone. Thyroglobulin is contained in a fluid called colloid, and TSH stimulation results in higher levels of colloid accumulation in the thyroid. In the absence of iodine, this is not converted to thyroid hormone, and colloid begins to accumulate more and more in the thyroid gland, leading to goiter.
Disorders can arise from both the underproduction and overproduction of thyroid hormones. Hypothyroidism, underproduction of the thyroid hormones, can cause a low metabolic rate leading to weight gain, sensitivity to cold, and reduced mental activity, among other symptoms. In children, hypothyroidism can cause cretinism, which can lead to mental retardation and growth defects. Hyperthyroidism, the overproduction of thyroid hormones, can lead to an increased metabolic rate and its effects: weight loss, excess heat production, sweating, and an increased heart rate. Graves’ disease is one example of a hyperthyroid condition.
Hormonal Control of Blood Calcium Levels
Regulation of blood calcium concentrations is important for generation of muscle contractions and nerve impulses, which are electrically stimulated. If calcium levels get too high, membrane permeability to sodium decreases and membranes become less responsive. If calcium levels get too low, membrane permeability to sodium increases and convulsions or muscle spasms can result.
Blood calcium levels are regulated by parathyroid hormone (PTH), which is produced by the parathyroid glands, as illustrated in Figure \(6\). PTH is released in response to low blood Ca2+ levels. PTH increases Ca2+ levels by targeting the skeleton, the kidneys, and the intestine. In the skeleton, PTH stimulates osteoclasts, which causes bone to be reabsorbed, releasing Ca2+ from bone into the blood. PTH also inhibits osteoblasts, reducing Ca2+ deposition in bone. In the intestines, PTH increases dietary Ca2+ absorption, and in the kidneys, PTH stimulates reabsorption of the CA2+. While PTH acts directly on the kidneys to increase Ca2+ reabsorption, its effects on the intestine are indirect. PTH triggers the formation of calcitriol, an active form of vitamin D, which acts on the intestines to increase absorption of dietary calcium. PTH release is inhibited by rising blood calcium levels.
Hyperparathyroidism results from an overproduction of parathyroid hormone. This results in excessive calcium being removed from bones and introduced into blood circulation, producing structural weakness of the bones, which can lead to deformation and fractures, plus nervous system impairment due to high blood calcium levels. Hypoparathyroidism, the underproduction of PTH, results in extremely low levels of blood calcium, which causes impaired muscle function and may result in tetany (severe sustained muscle contraction).
The hormone calcitonin, which is produced by the parafollicular or C cells of the thyroid, has the opposite effect on blood calcium levels as does PTH. Calcitonin decreases blood calcium levels by inhibiting osteoclasts, stimulating osteoblasts, and stimulating calcium excretion by the kidneys. This results in calcium being added to the bones to promote structural integrity. Calcitonin is most important in children (when it stimulates bone growth), during pregnancy (when it reduces maternal bone loss), and during prolonged starvation (because it reduces bone mass loss). In healthy nonpregnant, unstarved adults, the role of calcitonin is unclear.
Hormonal Regulation of Growth
Hormonal regulation is required for the growth and replication of most cells in the body. Growth hormone (GH), produced by the anterior portion of the pituitary gland, accelerates the rate of protein synthesis, particularly in skeletal muscle and bones. Growth hormone has direct and indirect mechanisms of action. The first direct action of GH is stimulation of triglyceride breakdown (lipolysis) and release into the blood by adipocytes. This results in a switch by most tissues from utilizing glucose as an energy source to utilizing fatty acids. This process is called a glucose-sparing effect. In another direct mechanism, GH stimulates glycogen breakdown in the liver; the glycogen is then released into the blood as glucose. Blood glucose levels increase as most tissues are utilizing fatty acids instead of glucose for their energy needs. The GH mediated increase in blood glucose levels is called a diabetogenic effect because it is similar to the high blood glucose levels seen in diabetes mellitus.
The indirect mechanism of GH action is mediated by insulin-like growth factors (IGFs) or somatomedins, which are a family of growth-promoting proteins produced by the liver, which stimulates tissue growth. IGFs stimulate the uptake of amino acids from the blood, allowing the formation of new proteins, particularly in skeletal muscle cells, cartilage cells, and other target cells, as shown in Figure \(7\). This is especially important after a meal, when glucose and amino acid concentration levels are high in the blood. GH levels are regulated by two hormones produced by the hypothalamus. GH release is stimulated by growth hormone-releasing hormone (GHRH) and is inhibited by growth hormone-inhibiting hormone (GHIH), also called somatostatin.
A balanced production of growth hormone is critical for proper development. Underproduction of GH in adults does not appear to cause any abnormalities, but in children it can result in pituitary dwarfism, in which growth is reduced. Pituitary dwarfism is characterized by symmetric body formation. In some cases, individuals are under 30 inches in height. Oversecretion of growth hormone can lead to gigantism in children, causing excessive growth. In some documented cases, individuals can reach heights of over eight feet. In adults, excessive GH can lead to acromegaly, a condition in which there is enlargement of bones in the face, hands, and feet that are still capable of growth.
Hormonal Regulation of Stress
When a threat or danger is perceived, the body responds by releasing hormones that will ready it for the “fight-or-flight” response. The effects of this response are familiar to anyone who has been in a stressful situation: increased heart rate, dry mouth, and hair standing up.
Evolution Connection: Fight-or-Flight Response
Interactions of the endocrine hormones have evolved to ensure the body’s internal environment remains stable. Stressors are stimuli that disrupt homeostasis. The sympathetic division of the vertebrate autonomic nervous system has evolved the fight-or-flight response to counter stress-induced disruptions of homeostasis. In the initial alarm phase, the sympathetic nervous system stimulates an increase in energy levels through increased blood glucose levels. This prepares the body for physical activity that may be required to respond to stress: to either fight for survival or to flee from danger.
However, some stresses, such as illness or injury, can last for a long time. Glycogen reserves, which provide energy in the short-term response to stress, are exhausted after several hours and cannot meet long-term energy needs. If glycogen reserves were the only energy source available, neural functioning could not be maintained once the reserves became depleted due to the nervous system’s high requirement for glucose. In this situation, the body has evolved a response to counter long-term stress through the actions of the glucocorticoids, which ensure that long-term energy requirements can be met. The glucocorticoids mobilize lipid and protein reserves, stimulate gluconeogenesis, conserve glucose for use by neural tissue, and stimulate the conservation of salts and water. The mechanisms to maintain homeostasis that are described here are those observed in the human body. However, the fight-or-flight response exists in some form in all vertebrates.
The sympathetic nervous system regulates the stress response via the hypothalamus. Stressful stimuli cause the hypothalamus to signal the adrenal medulla (which mediates short-term stress responses) via nerve impulses, and the adrenal cortex, which mediates long-term stress responses, via the hormone adrenocorticotropic hormone (ACTH), which is produced by the anterior pituitary.
Short-term Stress Response
When presented with a stressful situation, the body responds by calling for the release of hormones that provide a burst of energy. The hormones epinephrine (also known as adrenaline) and norepinephrine (also known as noradrenaline) are released by the adrenal medulla. How do these hormones provide a burst of energy? Epinephrine and norepinephrine increase blood glucose levels by stimulating the liver and skeletal muscles to break down glycogen and by stimulating glucose release by liver cells. Additionally, these hormones increase oxygen availability to cells by increasing the heart rate and dilating the bronchioles. The hormones also prioritize body function by increasing blood supply to essential organs such as the heart, brain, and skeletal muscles, while restricting blood flow to organs not in immediate need, such as the skin, digestive system, and kidneys. Epinephrine and norepinephrine are collectively called catecholamines.
Link to Learning
Watch this Discovery Channel animation describing the flight-or-flight response.
Long-term Stress Response
Long-term stress response differs from short-term stress response. The body cannot sustain the bursts of energy mediated by epinephrine and norepinephrine for long times. Instead, other hormones come into play. In a long-term stress response, the hypothalamus triggers the release of ACTH from the anterior pituitary gland. The adrenal cortex is stimulated by ACTH to release steroid hormones called corticosteroids. Corticosteroids turn on transcription of certain genes in the nuclei of target cells. They change enzyme concentrations in the cytoplasm and affect cellular metabolism. There are two main corticosteroids: glucocorticoids such as cortisol, and mineralocorticoids such as aldosterone. These hormones target the breakdown of fat into fatty acids in the adipose tissue. The fatty acids are released into the bloodstream for other tissues to use for ATP production. The glucocorticoids primarily affect glucose metabolism by stimulating glucose synthesis. Glucocorticoids also have anti-inflammatory properties through inhibition of the immune system. For example, cortisone is used as an anti-inflammatory medication; however, it cannot be used long term as it increases susceptibility to disease due to its immune-suppressing effects.
Mineralocorticoids function to regulate ion and water balance of the body. The hormone aldosterone stimulates the reabsorption of water and sodium ions in the kidney, which results in increased blood pressure and volume.
Hypersecretion of glucocorticoids can cause a condition known as Cushing’s disease, characterized by a shifting of fat storage areas of the body. This can cause the accumulation of adipose tissue in the face and neck, and excessive glucose in the blood. Hyposecretion of the corticosteroids can cause Addison’s disease, which may result in bronzing of the skin, hypoglycemia, and low electrolyte levels in the blood.
Summary
Water levels in the body are controlled by antidiuretic hormone (ADH), which is produced in the hypothalamus and triggers the reabsorption of water by the kidneys. Underproduction of ADH can cause diabetes insipidus. Aldosterone, a hormone produced by the adrenal cortex of the kidneys, enhances Na+ reabsorption from the extracellular fluids and subsequent water reabsorption by diffusion. The renin-angiotensin-aldosterone system is one way that aldosterone release is controlled.
The reproductive system is controlled by the gonadotropins follicle-stimulating hormone (FSH) and luteinizing hormone (LH), which are produced by the pituitary gland. Gonadotropin release is controlled by the hypothalamic hormone gonadotropin-releasing hormone (GnRH). FSH stimulates the maturation of sperm cells in males and is inhibited by the hormone inhibin, while LH stimulates the production of the androgen testosterone. FSH stimulates egg maturation in females, while LH stimulates the production of estrogens and progesterone. Estrogens are a group of steroid hormones produced by the ovaries that trigger the development of secondary sex characteristics in females as well as control the maturation of the ova. In females, the pituitary also produces prolactin, which stimulates milk production after childbirth, and oxytocin, which stimulates uterine contraction during childbirth and milk let-down during suckling.
Insulin is produced by the pancreas in response to rising blood glucose levels and allows cells to utilize blood glucose and store excess glucose for later use. Diabetes mellitus is caused by reduced insulin activity and causes high blood glucose levels, or hyperglycemia. Glucagon is released by the pancreas in response to low blood glucose levels and stimulates the breakdown of glycogen into glucose, which can be used by the body. The body’s basal metabolic rate is controlled by the thyroid hormones thyroxine (T4) and triiodothyronine (T3). The anterior pituitary produces thyroid stimulating hormone (TSH), which controls the release of T3 and T4 from the thyroid gland. Iodine is necessary in the production of thyroid hormone, and the lack of iodine can lead to a condition called goiter.
Parathyroid hormone (PTH) is produced by the parathyroid glands in response to low blood Ca2+ levels. The parafollicular cells of the thyroid produce calcitonin, which reduces blood Ca2+ levels. Growth hormone (GH) is produced by the anterior pituitary and controls the growth rate of muscle and bone. GH action is indirectly mediated by insulin-like growth factors (IGFs). Short-term stress causes the hypothalamus to trigger the adrenal medulla to release epinephrine and norepinephrine, which trigger the fight or flight response. Long-term stress causes the hypothalamus to trigger the anterior pituitary to release adrenocorticotropic hormone (ACTH), which causes the release of corticosteroids, glucocorticoids, and mineralocorticoids, from the adrenal cortex.
Glossary
acromegaly
condition caused by overproduction of GH in adults
Addison’s disease
disorder caused by the hyposecretion of corticosteroids
adrenocorticotropic hormone (ACTH)
hormone released by the anterior pituitary, which stimulates the adrenal cortex to release corticosteroids during the long-term stress response
aldosterone
steroid hormone produced by the adrenal cortex that stimulates the reabsorption of Na+ from extracellular fluids and secretion of K+.
androgen
male sex hormone such as testosterone
antidiuretic hormone (ADH)
hormone produced by the hypothalamus and released by the posterior pituitary that increases water reabsorption by the kidneys
calcitonin
hormone produced by the parafollicular cells of the thyroid gland that functions to lower blood Ca2+ levels and promote bone growth
corticosteroid
hormone released by the adrenal cortex in response to long-term stress
cortisol
glucocorticoid produced in response to stress
Cushing’s disease
disorder caused by the hypersecretion of glucocorticoids
diabetes insipidus
disorder caused by underproduction of ADH
diabetes mellitus
disorder caused by low levels of insulin activity
diabetogenic effect
effect of GH that causes blood glucose levels to rise similar to diabetes mellitus
epinephrine
hormone released by the adrenal medulla in response to a short term stress
estrogens
- a group of steroid hormones, including estradiol and several others, that are produced by the ovaries and elicit secondary sex characteristics in females as well as control the maturation of the ova
follicle-stimulating hormone (FSH)
hormone produced by the anterior pituitary that stimulates gamete production
gigantism
condition caused by overproduction of GH in children
glucagon
hormone produced by the alpha cells of the pancreas in response to low blood sugar; functions to raise blood sugar levels
glucocorticoid
corticosteroid that affects glucose metabolism
gluconeogenesis
synthesis of glucose from amino acids
glucose-sparing effect
effect of GH that causes tissues to use fatty acids instead of glucose as an energy source
glycogenolysis
breakdown of glycogen into glucose
goiter
enlargement of the thyroid gland caused by insufficient dietary iodine levels
gonadotropin
hormone that regulates the gonads, including FSH and LH
growth hormone (GH)
hormone produced by the anterior pituitary that promotes protein synthesis and body growth
growth hormone-inhibiting hormone (GHIH)
hormone produced by the hypothalamus that inhibits growth hormone production, also called somatostatin
growth hormone-releasing hormone (GHRH)
hormone released by the hypothalamus that triggers the release of GH
hyperglycemia
high blood sugar level
hyperthyroidism
overactivity of the thyroid gland
hypoglycemia
low blood sugar level
hypothyroidism
underactivity of the thyroid gland
insulin
hormone produced by the beta cells of the pancreas in response to high blood glucose levels; functions to lower blood glucose levels
insulin-like growth factor (IGF)
growth-promoting protein produced by the liver
mineralocorticoid
corticosteroid that affects ion and water balance
norepinephrine
hormone released by the adrenal medulla in response to a short-term stress hormone production by the gonads
osmoreceptor
receptor in the hypothalamus that monitors the concentration of electrolytes in the blood
oxytocin
hormone released by the posterior pituitary to stimulate uterine contractions during childbirth and milk let-down in the mammary glands
parathyroid hormone (PTH)
hormone produced by the parathyroid glands in response to low blood Ca2+ levels; functions to raise blood Ca2+ levels
pituitary dwarfism
condition caused by underproduction of GH in children
prolactin (PRL)
hormone produced by the anterior pituitary that stimulates milk production
prolactin-inhibiting hormone
hormone produced by the hypothalamus that inhibits the release of prolactin
prolactin-releasing hormone
hormone produced by the hypothalamus that stimulates the release of prolactin
renin
enzyme produced by the juxtaglomerular apparatus of the kidneys that reacts with angiotensinogen to cause the release of aldosterone
thyroglobulin
glycoprotein found in the thyroid that is converted into thyroid hormone
thyroid-stimulating hormone (TSH)
hormone produced by the anterior pituitary that controls the release of T3 and T4 from the thyroid gland
thyroxine (tetraiodothyronine, T4)
thyroid hormone that controls the basal metabolic rate
triiodothyronine (T3)
thyroid hormone that controls the basal metabolic rate | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.01%3A_Regulation_of_Body_Processes_by_Chemical_Messengers/44.1.02%3A_Regulation_of_Body_Processes.txt |
Skills to Develop
• Describe the role of different glands in the endocrine system
• Explain how the different glands work together to maintain homeostasis
Both the endocrine and nervous systems use chemical signals to communicate and regulate the body's physiology. The endocrine system releases hormones that act on target cells to regulate development, growth, energy metabolism, reproduction, and many behaviors. The nervous system releases neurotransmitters or neurohormones that regulate neurons, muscle cells, and endocrine cells. Because the neurons can regulate the release of hormones, the nervous and endocrine systems work in a coordinated manner to regulate the body's physiology.
Hypothalamic-Pituitary Axis
The hypothalamus in vertebrates integrates the endocrine and nervous systems. The hypothalamus is an endocrine organ located in the diencephalon of the brain. It receives input from the body and other brain areas and initiates endocrine responses to environmental changes. The hypothalamus acts as an endocrine organ, synthesizing hormones and transporting them along axons to the posterior pituitary gland. It synthesizes and secretes regulatory hormones that control the endocrine cells in the anterior pituitary gland. The hypothalamus contains autonomic centers that control endocrine cells in the adrenal medulla via neuronal control.
The pituitary gland, sometimes called the hypophysis or “master gland” is located at the base of the brain in the sella turcica, a groove of the sphenoid bone of the skull, illustrated in Figure \(1\). It is attached to the hypothalamus via a stalk called the pituitary stalk (or infundibulum). The anterior portion of the pituitary gland is regulated by releasing or release-inhibiting hormones produced by the hypothalamus, and the posterior pituitary receives signals via neurosecretory cells to release hormones produced by the hypothalamus. The pituitary has two distinct regions—the anterior pituitary and the posterior pituitary—which between them secrete nine different peptide or protein hormones. The posterior lobe of the pituitary gland contains axons of the hypothalamic neurons.
Anterior Pituitary
The anterior pituitary gland, or adenohypophysis, is surrounded by a capillary network that extends from the hypothalamus, down along the infundibulum, and to the anterior pituitary. This capillary network is a part of the hypophyseal portal system that carries substances from the hypothalamus to the anterior pituitary and hormones from the anterior pituitary into the circulatory system. A portal system carries blood from one capillary network to another; therefore, the hypophyseal portal system allows hormones produced by the hypothalamus to be carried directly to the anterior pituitary without first entering the circulatory system.
The anterior pituitary produces seven hormones: growth hormone (GH), prolactin (PRL), thyroid-stimulating hormone (TSH), melanin-stimulating hormone (MSH), adrenocorticotropic hormone (ACTH), follicle-stimulating hormone (FSH), and luteinizing hormone (LH). Anterior pituitary hormones are sometimes referred to as tropic hormones, because they control the functioning of other organs. While these hormones are produced by the anterior pituitary, their production is controlled by regulatory hormones produced by the hypothalamus. These regulatory hormones can be releasing hormones or inhibiting hormones, causing more or less of the anterior pituitary hormones to be secreted. These travel from the hypothalamus through the hypophyseal portal system to the anterior pituitary where they exert their effect. Negative feedback then regulates how much of these regulatory hormones are released and how much anterior pituitary hormone is secreted.
Posterior Pituitary
The posterior pituitary is significantly different in structure from the anterior pituitary. It is a part of the brain, extending down from the hypothalamus, and contains mostly nerve fibers and neuroglial cells, which support axons that extend from the hypothalamus to the posterior pituitary. The posterior pituitary and the infundibulum together are referred to as the neurohypophysis.
The hormones antidiuretic hormone (ADH), also known as vasopressin, and oxytocin are produced by neurons in the hypothalamus and transported within these axons along the infundibulum to the posterior pituitary. They are released into the circulatory system via neural signaling from the hypothalamus. These hormones are considered to be posterior pituitary hormones, even though they are produced by the hypothalamus, because that is where they are released into the circulatory system. The posterior pituitary itself does not produce hormones, but instead stores hormones produced by the hypothalamus and releases them into the blood stream.
Thyroid Gland
The thyroid gland is located in the neck, just below the larynx and in front of the trachea, as shown in Figure \(2\). It is a butterfly-shaped gland with two lobes that are connected by the isthmus. It has a dark red color due to its extensive vascular system. When the thyroid swells due to dysfunction, it can be felt under the skin of the neck.
The thyroid gland is made up of many spherical thyroid follicles, which are lined with a simple cuboidal epithelium. These follicles contain a viscous fluid, called colloid, which stores the glycoprotein thyroglobulin, the precursor to the thyroid hormones. The follicles produce hormones that can be stored in the colloid or released into the surrounding capillary network for transport to the rest of the body via the circulatory system.
Thyroid follicle cells synthesize the hormone thyroxine, which is also known as T4 because it contains four atoms of iodine, and triiodothyronine, also known as T3 because it contains three atoms of iodine. Follicle cells are stimulated to release stored T3 and T4 by thyroid stimulating hormone (TSH), which is produced by the anterior pituitary. These thyroid hormones increase the rates of mitochondrial ATP production.
A third hormone, calcitonin, is produced by parafollicular cells of the thyroid either releasing hormones or inhibiting hormones. Calcitonin release is not controlled by TSH, but instead is released when calcium ion concentrations in the blood rise. Calcitonin functions to help regulate calcium concentrations in body fluids. It acts in the bones to inhibit osteoclast activity and in the kidneys to stimulate excretion of calcium. The combination of these two events lowers body fluid levels of calcium.
Parathyroid Glands
Most people have four parathyroid glands; however, the number can vary from two to six. These glands are located on the posterior surface of the thyroid gland, as shown in Figure \(3\). Normally, there is a superior gland and an inferior gland associated with each of the thyroid’s two lobes. Each parathyroid gland is covered by connective tissue and contains many secretory cells that are associated with a capillary network.
The parathyroid glands produce parathyroid hormone (PTH). PTH increases blood calcium concentrations when calcium ion levels fall below normal. PTH (1) enhances reabsorption of Ca2+ by the kidneys, (2) stimulates osteoclast activity and inhibits osteoblast activity, and (3) it stimulates synthesis and secretion of calcitriol by the kidneys, which enhances Ca2+ absorption by the digestive system. PTH is produced by chief cells of the parathyroid. PTH and calcitonin work in opposition to one another to maintain homeostatic Ca2+ levels in body fluids. Another type of cells, oxyphil cells, exist in the parathyroid but their function is not known. These hormones encourage bone growth, muscle mass, and blood cell formation in children and women.
Adrenal Glands
The adrenal glands are associated with the kidneys; one gland is located on top of each kidney as illustrated in Figure \(4\). The adrenal glands consist of an outer adrenal cortex and an inner adrenal medulla. These regions secrete different hormones.
Adrenal Cortex
The adrenal cortex is made up of layers of epithelial cells and associated capillary networks. These layers form three distinct regions: an outer zona glomerulosa that produces mineralocorticoids, a middle zona fasciculata that produces glucocorticoids, and an inner zona reticularis that produces androgens.
The main mineralocorticoid is aldosterone, which regulates the concentration of Na+ ions in urine, sweat, pancreas, and saliva. Aldosterone release from the adrenal cortex is stimulated by a decrease in blood concentrations of sodium ions, blood volume, or blood pressure, or by an increase in blood potassium levels.
The three main glucocorticoids are cortisol, corticosterone, and cortisone. The glucocorticoids stimulate the synthesis of glucose and gluconeogenesis (converting a non-carbohydrate to glucose) by liver cells and they promote the release of fatty acids from adipose tissue. These hormones increase blood glucose levels to maintain levels within a normal range between meals. These hormones are secreted in response to ACTH and levels are regulated by negative feedback.
Androgens are sex hormones that promote masculinity. They are produced in small amounts by the adrenal cortex in both males and females. They do not affect sexual characteristics and may supplement sex hormones released from the gonads.
Adrenal Medulla
The adrenal medulla contains large, irregularly shaped cells that are closely associated with blood vessels. These cells are innervated by preganglionic autonomic nerve fibers from the central nervous system.
The adrenal medulla contains two types of secretory cells: one that produces epinephrine (adrenaline) and another that produces norepinephrine (noradrenaline). Epinephrine is the primary adrenal medulla hormone accounting for 75 to 80 percent of its secretions. Epinephrine and norepinephrine increase heart rate, breathing rate, cardiac muscle contractions, blood pressure, and blood glucose levels. They also accelerate the breakdown of glucose in skeletal muscles and stored fats in adipose tissue.
The release of epinephrine and norepinephrine is stimulated by neural impulses from the sympathetic nervous system. Secretion of these hormones is stimulated by acetylcholine release from preganglionic sympathetic fibers innervating the adrenal medulla. These neural impulses originate from the hypothalamus in response to stress to prepare the body for the fight-or-flight response.
Pancreas
The pancreas, illustrated in Figure \(5\), is an elongated organ that is located between the stomach and the proximal portion of the small intestine. It contains both exocrine cells that excrete digestive enzymes and endocrine cells that release hormones. It is sometimes referred to as a heterocrine gland because it has both endocrine and exocrine functions.
The endocrine cells of the pancreas form clusters called pancreatic islets or the islets of Langerhans, as visible in the micrograph shown in Figure \(6\). The pancreatic islets contain two primary cell types: alpha cells, which produce the hormone glucagon, and beta cells, which produce the hormone insulin. These hormones regulate blood glucose levels. As blood glucose levels decline, alpha cells release glucagon to raise the blood glucose levels by increasing rates of glycogen breakdown and glucose release by the liver. When blood glucose levels rise, such as after a meal, beta cells release insulin to lower blood glucose levels by increasing the rate of glucose uptake in most body cells, and by increasing glycogen synthesis in skeletal muscles and the liver. Together, glucagon and insulin regulate blood glucose levels.
Pineal Gland
The pineal gland produces melatonin. The rate of melatonin production is affected by the photoperiod. Collaterals from the visual pathways innervate the pineal gland. During the day photoperiod, little melatonin is produced; however, melatonin production increases during the dark photoperiod (night). In some mammals, melatonin has an inhibitory affect on reproductive functions by decreasing production and maturation of sperm, oocytes, and reproductive organs. Melatonin is an effective antioxidant, protecting the CNS from free radicals such as nitric oxide and hydrogen peroxide. Lastly, melatonin is involved in biological rhythms, particularly circadian rhythms such as the sleep-wake cycle and eating habits.
Gonads
The gonads—the male testes and female ovaries—produce steroid hormones. The testes produce androgens, testosterone being the most prominent, which allow for the development of secondary sex characteristics and the production of sperm cells. The ovaries produce estradiol and progesterone, which cause secondary sex characteristics and prepare the body for childbirth.
Table \(1\): Endocrine Glands and their Associated Hormones
Endocrine Gland Associated Hormones Effect
Hypothalamus releasing and inhibiting hormones regulate hormone release from pituitary gland; produce oxytocin; produce uterine contractions and milk secretion in females
antidiuretic hormone (ADH) water reabsorption from kidneys; vasoconstriction to increase blood pressure
Pituitary (Anterior) growth hormone (GH) promotes growth of body tissues, protein synthesis; metabolic functions
prolactin (PRL) promotes milk production
thyroid stimulating hormone (TSH) stimulates thyroid hormone release
adrenocorticotropic hormone (ACTH) stimulates hormone release by adrenal cortex, glucocorticoids
follicle-stimulating hormone (FSH) stimulates gamete production (both ova and sperm); secretion of estradiol
luteinizing hormone (LH) stimulates androgen production by gonads; ovulation, secretion of progesterone
melanocyte-stimulating hormone (MSH) stimulates melanocytes of the skin increasing melanin pigment production.
Pituitary (Posterior) antidiuretic hormone (ADH) stimulates water reabsorption by kidneys
oxytocin stimulates uterine contractions during childbirth; milk ejection; stimulates ductus deferens and prostate gland contraction during emission
Thyroid thyroxine, triiodothyronine stimulate and maintain metabolism; growth and development
calcitonin reduces blood Ca2+ levels
Parathyroid parathyroid hormone (PTH) increases blood Ca2+ levels
Adrenal (Cortex) aldosterone increases blood Na+ levels; increase K+ secretion
cortisol, corticosterone, cortisone increase blood glucose levels; anti-inflammatory effects
Adrenal (Medulla) epinephrine, norepinephrine stimulate fight-or-flight response; increase blood gluclose levels; increase metabolic activities
Pancreas insulin reduces blood glucose levels
glucagon increases blood glucose levels
Pineal gland melatonin regulates some biological rhythms and protects CNS from free radicals
Testes androgens regulate, promote, increase or maintain sperm production; male secondary sexual characteristics
Ovaries estrogen promotes uterine lining growth; female secondary sexual characteristics
progestins promote and maintain uterine lining growth
Organs with Secondary Endocrine Functions
There are several organs whose primary functions are non-endocrine but that also possess endocrine functions. These include the heart, kidneys, intestines, thymus, gonads, and adipose tissue.
The heart possesses endocrine cells in the walls of the atria that are specialized cardiac muscle cells. These cells release the hormone atrial natriuretic peptide (ANP) in response to increased blood volume. High blood volume causes the cells to be stretched, resulting in hormone release. ANP acts on the kidneys to reduce the reabsorption of Na+, causing Na+ and water to be excreted in the urine. ANP also reduces the amounts of renin released by the kidneys and aldosterone released by the adrenal cortex, further preventing the retention of water. In this way, ANP causes a reduction in blood volume and blood pressure, and reduces the concentration of Na+ in the blood.
The gastrointestinal tract produces several hormones that aid in digestion. The endocrine cells are located in the mucosa of the GI tract throughout the stomach and small intestine. Some of the hormones produced include gastrin, secretin, and cholecystokinin, which are secreted in the presence of food, and some of which act on other organs such as the pancreas, gallbladder, and liver. They trigger the release of gastric juices, which help to break down and digest food in the GI tract.
While the adrenal glands associated with the kidneys are major endocrine glands, the kidneys themselves also possess endocrine function. Renin is released in response to decreased blood volume or pressure and is part of the renin-angiotensin-aldosterone system that leads to the release of aldosterone. Aldosterone then causes the retention of Na+ and water, raising blood volume. The kidneys also release calcitriol, which aids in the absorption of Ca2+ and phosphate ions. Erythropoietin (EPO) is a protein hormone that triggers the formation of red blood cells in the bone marrow. EPO is released in response to low oxygen levels. Because red blood cells are oxygen carriers, increased production results in greater oxygen delivery throughout the body. EPO has been used by athletes to improve performance, as greater oxygen delivery to muscle cells allows for greater endurance. Because red blood cells increase the viscosity of blood, artificially high levels of EPO can cause severe health risks.
The thymus is found behind the sternum; it is most prominent in infants, becoming smaller in size through adulthood. The thymus produces hormones referred to as thymosins, which contribute to the development of the immune response.
Adipose tissue is a connective tissue found throughout the body. It produces the hormone leptin in response to food intake. Leptin increases the activity of anorexigenic neurons and decreases that of orexigenic neurons, producing a feeling of satiety after eating, thus affecting appetite and reducing the urge for further eating. Leptin is also associated with reproduction. It must be present for GnRH and gonadotropin synthesis to occur. Extremely thin females may enter puberty late; however, if adipose levels increase, more leptin will be produced, improving fertility.
Summary
The pituitary gland is located at the base of the brain and is attached to the hypothalamus by the infundibulum. The anterior pituitary receives products from the hypothalamus by the hypophyseal portal system and produces six hormones. The posterior pituitary is an extension of the brain and releases hormones (antidiuretic hormone and oxytocin) produced by the hypothalamus.
The thyroid gland is located in the neck and is composed of two lobes connected by the isthmus. The thyroid is made up of follicle cells that produce the hormones thyroxine and triiodothyronine. Parafollicular cells of the thyroid produce calcitonin. The parathyroid glands lie on the posterior surface of the thyroid gland and produce parathyroid hormone.
The adrenal glands are located on top of the kidneys and consist of the renal cortex and renal medulla. The adrenal cortex is the outer part of the adrenal gland and produces the corticosteroids, glucocorticoids, and mineralocorticoids. The adrenal medulla is the inner part of the adrenal gland and produces the catecholamines epinephrine and norepinephrine.
The pancreas lies in the abdomen between the stomach and the small intestine. Clusters of endocrine cells in the pancreas form the islets of Langerhans, which are composed of alpha cells that release glucagon and beta cells that release insulin.
Some organs possess endocrine activity as a secondary function but have another primary function. The heart produces the hormone atrial natriuretic peptide, which functions to reduce blood volume, pressure, and Na+ concentration. The gastrointestinal tract produces various hormones that aid in digestion. The kidneys produce renin, calcitriol, and erythropoietin. Adipose tissue produces leptin, which promotes satiety signals in the brain.
Glossary
adrenal cortex
outer portion of adrenal glands that produces corticosteroids
adrenal gland
endocrine glands associated with the kidneys
adrenal medulla
inner portion of adrenal glands that produces epinephrine and norepinephrine
alpha cell
endocrine cell of the pancreatic islets that produces the hormone glucagon
anterior pituitary
portion of the pituitary gland that produces six hormones; also called adenohypophysis
atrial natriuretic peptide (ANP)
hormone produced by the heart to reduce blood volume, pressure, and Na+ concentration
beta cell
endocrine cell of the pancreatic islets that produces the hormone insulin
colloid
fluid inside the thyroid gland that contains the glycoprotein thyroglobulin
endocrine gland
gland that secretes hormones into the surrounding interstitial fluid, which then diffuse into blood and are carried to various organs and tissues within the body
erythropoietin (EPO)
hormone produced by the kidneys to stimulate red blood cell production in the bone marrow
hypophyseal portal system
system of blood vessels that carries hormones from the hypothalamus to the anterior pituitary
islets of Langerhans (pancreatic islets)
endocrine cells of the pancreas
isthmus
tissue mass that connects the two lobes of the thyroid gland
leptin
hormone produced by adipose tissue that promotes feelings of satiety and reduces hunger
pancreas
organ located between the stomach and the small intestine that contains exocrine and endocrine cells
parafollicular cell
thyroid cell that produces the hormone calcitonin
parathyroid gland
gland located on the surface of the thyroid that produces parathyroid hormone
pituitary gland
endocrine gland located at the base of the brain composed of an anterior and posterior region; also called hypophysis
pituitary stalk
(also, infundibulum) stalk that connects the pituitary gland to the hypothalamus
posterior pituitary
extension of the brain that releases hormones produced by the hypothalamus; along with the infundibulum, it is also referred to as the neurohypophysis
thymus
gland located behind the sternum that produces thymosin hormones that contribute to the development of the immune system
thyroid gland
endocrine gland located in the neck that produces thyroid hormones thyroxine and triiodothyronine | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.01%3A_Regulation_of_Body_Processes_by_Chemical_Messengers/44.1.03%3A_Endocrine_Glands.txt |
Skills to Develop
• Explain how hormones work
• Discuss the role of different types of hormone receptors
Hormones mediate changes in target cells by binding to specific hormone receptors. In this way, even though hormones circulate throughout the body and come into contact with many different cell types, they only affect cells that possess the necessary receptors. Receptors for a specific hormone may be found on many different cells or may be limited to a small number of specialized cells. For example, thyroid hormones act on many different tissue types, stimulating metabolic activity throughout the body. Cells can have many receptors for the same hormone but often also possess receptors for different types of hormones. The number of receptors that respond to a hormone determines the cell’s sensitivity to that hormone, and the resulting cellular response. Additionally, the number of receptors that respond to a hormone can change over time, resulting in increased or decreased cell sensitivity. In up-regulation, the number of receptors increases in response to rising hormone levels, making the cell more sensitive to the hormone and allowing for more cellular activity. When the number of receptors decreases in response to rising hormone levels, called down-regulation, cellular activity is reduced.
Receptor binding alters cellular activity and results in an increase or decrease in normal body processes. Depending on the location of the protein receptor on the target cell and the chemical structure of the hormone, hormones can mediate changes directly by binding to intracellular hormone receptors and modulating gene transcription, or indirectly by binding to cell surface receptors and stimulating signaling pathways.
Intracellular Hormone Receptors
Lipid-derived (soluble) hormones such as steroid hormones diffuse across the membranes of the endocrine cell. Once outside the cell, they bind to transport proteins that keep them soluble in the bloodstream. At the target cell, the hormones are released from the carrier protein and diffuse across the lipid bilayer of the plasma membrane of cells. The steroid hormones pass through the plasma membrane of a target cell and adhere to intracellular receptors residing in the cytoplasm or in the nucleus. The cell signaling pathways induced by the steroid hormones regulate specific genes on the cell's DNA. The hormones and receptor complex act as transcription regulators by increasing or decreasing the synthesis of mRNA molecules of specific genes. This, in turn, determines the amount of corresponding protein that is synthesized by altering gene expression. This protein can be used either to change the structure of the cell or to produce enzymes that catalyze chemical reactions. In this way, the steroid hormone regulates specific cell processes as illustrated in Figure \(1\).
Exercise \(1\)
Heat shock proteins (HSP) are so named because they help refold misfolded proteins. In response to increased temperature (a “heat shock”), heat shock proteins are activated by release from the NR/HSP complex. At the same time, transcription of HSP genes is activated. Why do you think the cell responds to a heat shock by increasing the activity of proteins that help refold misfolded proteins?
Answer
Proteins unfold, or denature, at higher temperatures.
Other lipid-soluble hormones that are not steroid hormones, such as vitamin D and thyroxine, have receptors located in the nucleus. The hormones diffuse across both the plasma membrane and the nuclear envelope, then bind to receptors in the nucleus. The hormone-receptor complex stimulates transcription of specific genes.
Plasma Membrane Hormone Receptors
Amino acid derived hormones and polypeptide hormones are not lipid-derived (lipid-soluble) and therefore cannot diffuse through the plasma membrane of cells. Lipid insoluble hormones bind to receptors on the outer surface of the plasma membrane, via plasma membrane hormone receptors. Unlike steroid hormones, lipid insoluble hormones do not directly affect the target cell because they cannot enter the cell and act directly on DNA. Binding of these hormones to a cell surface receptor results in activation of a signaling pathway; this triggers intracellular activity and carries out the specific effects associated with the hormone. In this way, nothing passes through the cell membrane; the hormone that binds at the surface remains at the surface of the cell while the intracellular product remains inside the cell. The hormone that initiates the signaling pathway is called a first messenger, which activates a second messenger in the cytoplasm, as illustrated in Figure \(2\).
One very important second messenger is cyclic AMP (cAMP). When a hormone binds to its membrane receptor, a G-protein that is associated with the receptor is activated; G-proteins are proteins separate from receptors that are found in the cell membrane. When a hormone is not bound to the receptor, the G-protein is inactive and is bound to guanosine diphosphate, or GDP. When a hormone binds to the receptor, the G-protein is activated by binding guanosine triphosphate, or GTP, in place of GDP. After binding, GTP is hydrolysed by the G-protein into GDP and becomes inactive.
The activated G-protein in turn activates a membrane-bound enzyme called adenylyl cyclase. Adenylyl cyclase catalyzes the conversion of ATP to cAMP. cAMP, in turn, activates a group of proteins called protein kinases, which transfer a phosphate group from ATP to a substrate molecule in a process called phosphorylation. The phosphorylation of a substrate molecule changes its structural orientation, thereby activating it. These activated molecules can then mediate changes in cellular processes.
The effect of a hormone is amplified as the signaling pathway progresses. The binding of a hormone at a single receptor causes the activation of many G-proteins, which activates adenylyl cyclase. Each molecule of adenylyl cyclase then triggers the formation of many molecules of cAMP. Further amplification occurs as protein kinases, once activated by cAMP, can catalyze many reactions. In this way, a small amount of hormone can trigger the formation of a large amount of cellular product. To stop hormone activity, cAMP is deactivated by the cytoplasmic enzyme phosphodiesterase, or PDE. PDE is always present in the cell and breaks down cAMP to control hormone activity, preventing overproduction of cellular products.
The specific response of a cell to a lipid insoluble hormone depends on the type of receptors that are present on the cell membrane and the substrate molecules present in the cell cytoplasm. Cellular responses to hormone binding of a receptor include altering membrane permeability and metabolic pathways, stimulating synthesis of proteins and enzymes, and activating hormone release.
Summary
Hormones cause cellular changes by binding to receptors on target cells. The number of receptors on a target cell can increase or decrease in response to hormone activity. Hormones can affect cells directly through intracellular hormone receptors or indirectly through plasma membrane hormone receptors.
Lipid-derived (soluble) hormones can enter the cell by diffusing across the plasma membrane and binding to DNA to regulate gene transcription and to change the cell’s activities by inducing production of proteins that affect, in general, the long-term structure and function of the cell. Lipid insoluble hormones bind to receptors on the plasma membrane surface and trigger a signaling pathway to change the cell’s activities by inducing production of various cell products that affect the cell in the short-term. The hormone is called a first messenger and the cellular component is called a second messenger. G-proteins activate the second messenger (cyclic AMP), triggering the cellular response. Response to hormone binding is amplified as the signaling pathway progresses. Cellular responses to hormones include the production of proteins and enzymes and altered membrane permeability.
Glossary
adenylate cyclase
an enzyme that catalyzes the conversion of ATP to cyclic AMP
down-regulation
a decrease in the number of hormone receptors in response to increased hormone levels
first messenger
the hormone that binds to a plasma membrane hormone receptor to trigger a signal transduction pathway
G-protein
a membrane protein activated by the hormone first messenger to activate formation of cyclic AMP
hormone receptor
the cellular protein that binds to a hormone
intracellular hormone receptor
a hormone receptor in the cytoplasm or nucleus of a cell
phosphodiesterase (PDE)
enzyme that deactivates cAMP, stopping hormone activity
plasma membrane hormone receptor
a hormone receptor on the surface of the plasma membrane of a cell
up-regulation
an increase in the number of hormone receptors in response to increased hormone levels | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.02%3A_Overview_of_Hormone_Action.txt |
Skills to Develop
• List the different types of hormones
• Explain their role in maintaining homeostasis
Maintaining homeostasis within the body requires the coordination of many different systems and organs. Communication between neighboring cells, and between cells and tissues in distant parts of the body, occurs through the release of chemicals called hormones. Hormones are released into body fluids (usually blood) that carry these chemicals to their target cells. At the target cells, which are cells that have a receptor for a signal or ligand from a signal cell, the hormones elicit a response. The cells, tissues, and organs that secrete hormones make up the endocrine system. Examples of glands of the endocrine system include the adrenal glands, which produce hormones such as epinephrine and norepinephrine that regulate responses to stress, and the thyroid gland, which produces thyroid hormones that regulate metabolic rates.
Although there are many different hormones in the human body, they can be divided into three classes based on their chemical structure: lipid-derived, amino acid-derived, and peptide (peptide and proteins) hormones. One of the key distinguishing features of lipid-derived hormones is that they can diffuse across plasma membranes whereas the amino acid-derived and peptide hormones cannot.
Lipid-Derived Hormones (or Lipid-soluble Hormones)
Most lipid hormones are derived from cholesterol and thus are structurally similar to it, as illustrated in Figure \(1\). The primary class of lipid hormones in humans is the steroid hormones. Chemically, these hormones are usually ketones or alcohols; their chemical names will end in “-ol” for alcohols or “-one” for ketones. Examples of steroid hormones include estradiol, which is an estrogen, or female sex hormone, and testosterone, which is an androgen, or male sex hormone. These two hormones are released by the female and male reproductive organs, respectively. Other steroid hormones include aldosterone and cortisol, which are released by the adrenal glands along with some other types of androgens. Steroid hormones are insoluble in water, and they are transported by transport proteins in blood. As a result, they remain in circulation longer than peptide hormones. For example, cortisol has a half-life of 60 to 90 minutes, while epinephrine, an amino acid derived-hormone, has a half-life of approximately one minute.
Amino Acid-Derived Hormones
The amino acid-derived hormones are relatively small molecules that are derived from the amino acids tyrosine and tryptophan, shown in Figure \(2\). If a hormone is amino acid-derived, its chemical name will end in “-ine”. Examples of amino acid-derived hormones include epinephrine and norepinephrine, which are synthesized in the medulla of the adrenal glands, and thyroxine, which is produced by the thyroid gland. The pineal gland in the brain makes and secretes melatonin which regulates sleep cycles.
Peptide Hormones
The structure of peptide hormones is that of a polypeptide chain (chain of amino acids). The peptide hormones include molecules that are short polypeptide chains, such as antidiuretic hormone and oxytocin produced in the brain and released into the blood in the posterior pituitary gland. This class also includes small proteins, like growth hormones produced by the pituitary, and large glycoproteins such as follicle-stimulating hormone produced by the pituitary. Figure \(3\) illustrates these peptide hormones.
Secreted peptides like insulin are stored within vesicles in the cells that synthesize them. They are then released in response to stimuli such as high blood glucose levels in the case of insulin. Amino acid-derived and polypeptide hormones are water-soluble and insoluble in lipids. These hormones cannot pass through plasma membranes of cells; therefore, their receptors are found on the surface of the target cells.
Career Connection: Endocrinologist
An endocrinologist is a medical doctor who specializes in treating disorders of the endocrine glands, hormone systems, and glucose and lipid metabolic pathways. An endocrine surgeon specializes in the surgical treatment of endocrine diseases and glands. Some of the diseases that are managed by endocrinologists: disorders of the pancreas (diabetes mellitus), disorders of the pituitary (gigantism, acromegaly, and pituitary dwarfism), disorders of the thyroid gland (goiter and Graves’ disease), and disorders of the adrenal glands (Cushing’s disease and Addison’s disease).
Endocrinologists are required to assess patients and diagnose endocrine disorders through extensive use of laboratory tests. Many endocrine diseases are diagnosed using tests that stimulate or suppress endocrine organ functioning. Blood samples are then drawn to determine the effect of stimulating or suppressing an endocrine organ on the production of hormones. For example, to diagnose diabetes mellitus, patients are required to fast for 12 to 24 hours. They are then given a sugary drink, which stimulates the pancreas to produce insulin to decrease blood glucose levels. A blood sample is taken one to two hours after the sugar drink is consumed. If the pancreas is functioning properly, the blood glucose level will be within a normal range. Another example is the A1C test, which can be performed during blood screening. The A1C test measures average blood glucose levels over the past two to three months by examining how well the blood glucose is being managed over a long time.
Once a disease has been diagnosed, endocrinologists can prescribe lifestyle changes and/or medications to treat the disease. Some cases of diabetes mellitus can be managed by exercise, weight loss, and a healthy diet; in other cases, medications may be required to enhance insulin release. If the disease cannot be controlled by these means, the endocrinologist may prescribe insulin injections.
In addition to clinical practice, endocrinologists may also be involved in primary research and development activities. For example, ongoing islet transplant research is investigating how healthy pancreas islet cells may be transplanted into diabetic patients. Successful islet transplants may allow patients to stop taking insulin injections.
Summary
There are three basic types of hormones: lipid-derived, amino acid-derived, and peptide. Lipid-derived hormones are structurally similar to cholesterol and include steroid hormones such as estradiol and testosterone. Amino acid-derived hormones are relatively small molecules and include the adrenal hormones epinephrine and norepinephrine. Peptide hormones are polypeptide chains or proteins and include the pituitary hormones, antidiuretic hormone (vasopressin), and oxytocin.
Glossary
amino acid-derived hormone
hormone derived from amino acids
lipid-derived hormone
hormone derived mostly from cholesterol
peptide hormone
hormone composed of a polypeptide chain
44.03: The Pituitary and Hypothalamus- The Body's Control Centers
Skills to Develop
• Explain how hormones regulate the excretory system
• Discuss the role of hormones in the reproductive system
• Describe how hormones regulate metabolism
• Explain the role of hormones in different diseases
Hormones have a wide range of effects and modulate many different body processes. The key regulatory processes that will be examined here are those affecting the excretory system, the reproductive system, metabolism, blood calcium concentrations, growth, and the stress response.
Hormonal Regulation of the Excretory System
Maintaining a proper water balance in the body is important to avoid dehydration or over-hydration (hyponatremia). The water concentration of the body is monitored by osmoreceptors in the hypothalamus, which detect the concentration of electrolytes in the extracellular fluid. The concentration of electrolytes in the blood rises when there is water loss caused by excessive perspiration, inadequate water intake, or low blood volume due to blood loss. An increase in blood electrolyte levels results in a neuronal signal being sent from the osmoreceptors in hypothalamic nuclei. The pituitary gland has two components: anterior and posterior. The anterior pituitary is composed of glandular cells that secrete protein hormones. The posterior pituitary is an extension of the hypothalamus. It is composed largely of neurons that are continuous with the hypothalamus.
The hypothalamus produces a polypeptide hormone known as antidiuretic hormone (ADH), which is transported to and released from the posterior pituitary gland. The principal action of ADH is to regulate the amount of water excreted by the kidneys. As ADH (which is also known as vasopressin) causes direct water reabsorption from the kidney tubules, salts and wastes are concentrated in what will eventually be excreted as urine. The hypothalamus controls the mechanisms of ADH secretion, either by regulating blood volume or the concentration of water in the blood. Dehydration or physiological stress can cause an increase of osmolarity above 300 mOsm/L, which in turn, raises ADH secretion and water will be retained, causing an increase in blood pressure. ADH travels in the bloodstream to the kidneys. Once at the kidneys, ADH changes the kidneys to become more permeable to water by temporarily inserting water channels, aquaporins, into the kidney tubules. Water moves out of the kidney tubules through the aquaporins, reducing urine volume. The water is reabsorbed into the capillaries lowering blood osmolarity back toward normal. As blood osmolarity decreases, a negative feedback mechanism reduces osmoreceptor activity in the hypothalamus, and ADH secretion is reduced. ADH release can be reduced by certain substances, including alcohol, which can cause increased urine production and dehydration.
Chronic underproduction of ADH or a mutation in the ADH receptor results in diabetes insipidus. If the posterior pituitary does not release enough ADH, water cannot be retained by the kidneys and is lost as urine. This causes increased thirst, but water taken in is lost again and must be continually consumed. If the condition is not severe, dehydration may not occur, but severe cases can lead to electrolyte imbalances due to dehydration.
Another hormone responsible for maintaining electrolyte concentrations in extracellular fluids is aldosterone, a steroid hormone that is produced by the adrenal cortex. In contrast to ADH, which promotes the reabsorption of water to maintain proper water balance, aldosterone maintains proper water balance by enhancing Na+ reabsorption and K+ secretion from extracellular fluid of the cells in kidney tubules. Because it is produced in the cortex of the adrenal gland and affects the concentrations of minerals Na+ and K+, aldosterone is referred to as a mineralocorticoid, a corticosteroid that affects ion and water balance. Aldosterone release is stimulated by a decrease in blood sodium levels, blood volume, or blood pressure, or an increase in blood potassium levels. It also prevents the loss of Na+ from sweat, saliva, and gastric juice. The reabsorption of Na+ also results in the osmotic reabsorption of water, which alters blood volume and blood pressure.
Aldosterone production can be stimulated by low blood pressure, which triggers a sequence of chemical release, as illustrated in Figure \(1\). When blood pressure drops, the renin-angiotensin-aldosterone system (RAAS) is activated. Cells in the juxtaglomerular apparatus, which regulates the functions of the nephrons of the kidney, detect this and release renin. Renin, an enzyme, circulates in the blood and reacts with a plasma protein produced by the liver called angiotensinogen. When angiotensinogen is cleaved by renin, it produces angiotensin I, which is then converted into angiotensin II in the lungs. Angiotensin II functions as a hormone and then causes the release of the hormone aldosterone by the adrenal cortex, resulting in increased Na+ reabsorption, water retention, and an increase in blood pressure. Angiotensin II in addition to being a potent vasoconstrictor also causes an increase in ADH and increased thirst, both of which help to raise blood pressure.
Hormonal Regulation of the Reproductive System
Regulation of the reproductive system is a process that requires the action of hormones from the pituitary gland, the adrenal cortex, and the gonads. During puberty in both males and females, the hypothalamus produces gonadotropin-releasing hormone (GnRH), which stimulates the production and release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary gland. These hormones regulate the gonads (testes in males and ovaries in females) and therefore are called gonadotropins. In both males and females, FSH stimulates gamete production and LH stimulates production of hormones by the gonads. An increase in gonad hormone levels inhibits GnRH production through a negative feedback loop.
Regulation of the Male Reproductive System
In males, FSH stimulates the maturation of sperm cells. FSH production is inhibited by the hormone inhibin, which is released by the testes. LH stimulates production of the sex hormones (androgens) by the interstitial cells of the testes and therefore is also called interstitial cell-stimulating hormone. The most widely known androgen in males is testosterone. Testosterone promotes the production of sperm and masculine characteristics. The adrenal cortex also produces small amounts of testosterone precursor, although the role of this additional hormone production is not fully understood.
Everyday Connection: The Dangers of Synthetic Hormones
Some athletes attempt to boost their performance by using artificial hormones that enhance muscle performance. Anabolic steroids, a form of the male sex hormone testosterone, are one of the most widely known performance-enhancing drugs. Steroids are used to help build muscle mass. Other hormones that are used to enhance athletic performance include erythropoietin, which triggers the production of red blood cells, and human growth hormone, which can help in building muscle mass. Most performance enhancing drugs are illegal for non-medical purposes. They are also banned by national and international governing bodies including the International Olympic Committee, the U.S. Olympic Committee, the National Collegiate Athletic Association, the Major League Baseball, and the National Football League.
The side effects of synthetic hormones are often significant and non-reversible, and in some cases, fatal. Androgens produce several complications such as liver dysfunctions and liver tumors, prostate gland enlargement, difficulty urinating, premature closure of epiphyseal cartilages, testicular atrophy, infertility, and immune system depression. The physiological strain caused by these substances is often greater than what the body can handle, leading to unpredictable and dangerous effects and linking their use to heart attacks, strokes, and impaired cardiac function.
Regulation of the Female Reproductive System
In females, FSH stimulates development of egg cells, called ova, which develop in structures called follicles. Follicle cells produce the hormone inhibin, which inhibits FSH production. LH also plays a role in the development of ova, induction of ovulation, and stimulation of estradiol and progesterone production by the ovaries, as illustrated in Figure \(3\). Estradiol and progesterone are steroid hormones that prepare the body for pregnancy. Estradiol produces secondary sex characteristics in females, while both estradiol and progesterone regulate the menstrual cycle.
In addition to producing FSH and LH, the anterior portion of the pituitary gland also produces the hormone prolactin (PRL) in females. Prolactin stimulates the production of milk by the mammary glands following childbirth. Prolactin levels are regulated by the hypothalamic hormones prolactin-releasing hormone (PRH) and prolactin-inhibiting hormone (PIH), which is now known to be dopamine. PRH stimulates the release of prolactin and PIH inhibits it.
The posterior pituitary releases the hormone oxytocin, which stimulates uterine contractions during childbirth. The uterine smooth muscles are not very sensitive to oxytocin until late in pregnancy when the number of oxytocin receptors in the uterus peaks. Stretching of tissues in the uterus and cervix stimulates oxytocin release during childbirth. Contractions increase in intensity as blood levels of oxytocin rise via a positive feedback mechanism until the birth is complete. Oxytocin also stimulates the contraction of myoepithelial cells around the milk-producing mammary glands. As these cells contract, milk is forced from the secretory alveoli into milk ducts and is ejected from the breasts in milk ejection (“let-down”) reflex. Oxytocin release is stimulated by the suckling of an infant, which triggers the synthesis of oxytocin in the hypothalamus and its release into circulation at the posterior pituitary.
Hormonal Regulation of Metabolism
Blood glucose levels vary widely over the course of a day as periods of food consumption alternate with periods of fasting. Insulin and glucagon are the two hormones primarily responsible for maintaining homeostasis of blood glucose levels. Additional regulation is mediated by the thyroid hormones.
Regulation of Blood Glucose Levels by Insulin and Glucagon
Cells of the body require nutrients in order to function, and these nutrients are obtained through feeding. In order to manage nutrient intake, storing excess intake and utilizing reserves when necessary, the body uses hormones to moderate energy stores. Insulin is produced by the beta cells of the pancreas, which are stimulated to release insulin as blood glucose levels rise (for example, after a meal is consumed). Insulin lowers blood glucose levels by enhancing the rate of glucose uptake and utilization by target cells, which use glucose for ATP production. It also stimulates the liver to convert glucose to glycogen, which is then stored by cells for later use. Insulin also increases glucose transport into certain cells, such as muscle cells and the liver. This results from an insulin-mediated increase in the number of glucose transporter proteins in cell membranes, which remove glucose from circulation by facilitated diffusion. As insulin binds to its target cell via insulin receptors and signal transduction, it triggers the cell to incorporate glucose transport proteins into its membrane. This allows glucose to enter the cell, where it can be used as an energy source. However, this does not occur in all cells: some cells, including those in the kidneys and brain, can access glucose without the use of insulin. Insulin also stimulates the conversion of glucose to fat in adipocytes and the synthesis of proteins. These actions mediated by insulin cause blood glucose concentrations to fall, called a hypoglycemic “low sugar” effect, which inhibits further insulin release from beta cells through a negative feedback loop.
Link to Learning
This animation describes the role of insulin and the pancreas in diabetes.
Impaired insulin function can lead to a condition called diabetes mellitus, the main symptoms of which are illustrated in Figure \(4\). This can be caused by low levels of insulin production by the beta cells of the pancreas, or by reduced sensitivity of tissue cells to insulin. This prevents glucose from being absorbed by cells, causing high levels of blood glucose, or hyperglycemia (high sugar). High blood glucose levels make it difficult for the kidneys to recover all the glucose from nascent urine, resulting in glucose being lost in urine. High glucose levels also result in less water being reabsorbed by the kidneys, causing high amounts of urine to be produced; this may result in dehydration. Over time, high blood glucose levels can cause nerve damage to the eyes and peripheral body tissues, as well as damage to the kidneys and cardiovascular system. Oversecretion of insulin can cause hypoglycemia, low blood glucose levels. This causes insufficient glucose availability to cells, often leading to muscle weakness, and can sometimes cause unconsciousness or death if left untreated.
When blood glucose levels decline below normal levels, for example between meals or when glucose is utilized rapidly during exercise, the hormone glucagon is released from the alpha cells of the pancreas. Glucagon raises blood glucose levels, eliciting what is called a hyperglycemic effect, by stimulating the breakdown of glycogen to glucose in skeletal muscle cells and liver cells in a process called glycogenolysis. Glucose can then be utilized as energy by muscle cells and released into circulation by the liver cells. Glucagon also stimulates absorption of amino acids from the blood by the liver, which then converts them to glucose. This process of glucose synthesis is called gluconeogenesis. Glucagon also stimulates adipose cells to release fatty acids into the blood. These actions mediated by glucagon result in an increase in blood glucose levels to normal homeostatic levels. Rising blood glucose levels inhibit further glucagon release by the pancreas via a negative feedback mechanism. In this way, insulin and glucagon work together to maintain homeostatic glucose levels, as shown in Figure \(5\).
Exercise
Pancreatic tumors may cause excess secretion of glucagon. Type I diabetes results from the failure of the pancreas to produce insulin. Which of the following statement about these two conditions is true?
1. A pancreatic tumor and type I diabetes will have the opposite effects on blood sugar levels.
2. A pancreatic tumor and type I diabetes will both cause hyperglycemia.
3. A pancreatic tumor and type I diabetes will both cause hypoglycemia.
4. Both pancreatic tumors and type I diabetes result in the inability of cells to take up glucose.
Answer
B
Regulation of Blood Glucose Levels by Thyroid Hormones
The basal metabolic rate, which is the amount of calories required by the body at rest, is determined by two hormones produced by the thyroid gland: thyroxine, also known as tetraiodothyronine or T4, and triiodothyronine, also known as T3. These hormones affect nearly every cell in the body except for the adult brain, uterus, testes, blood cells, and spleen. They are transported across the plasma membrane of target cells and bind to receptors on the mitochondria resulting in increased ATP production. In the nucleus, T3 and T4 activate genes involved in energy production and glucose oxidation. This results in increased rates of metabolism and body heat production, which is known as the hormone’s calorigenic effect.
T3 and T4 release from the thyroid gland is stimulated by thyroid-stimulating hormone (TSH), which is produced by the anterior pituitary. TSH binding at the receptors of the follicle of the thyroid triggers the production of T3 and T4 from a glycoprotein called thyroglobulin. Thyroglobulin is present in the follicles of the thyroid, and is converted into thyroid hormones with the addition of iodine. Iodine is formed from iodide ions that are actively transported into the thyroid follicle from the bloodstream. A peroxidase enzyme then attaches the iodine to the tyrosine amino acid found in thyroglobulin. T3 has three iodine ions attached, while T4 has four iodine ions attached. T3 and T4 are then released into the bloodstream, with T4 being released in much greater amounts than T3. As T3 is more active than T4 and is responsible for most of the effects of thyroid hormones, tissues of the body convert T4 to T3 by the removal of an iodine ion. Most of the released T3 and T4 becomes attached to transport proteins in the bloodstream and is unable to cross the plasma membrane of cells. These protein-bound molecules are only released when blood levels of the unattached hormone begin to decline. In this way, a week’s worth of reserve hormone is maintained in the blood. Increased T3 and T4 levels in the blood inhibit the release of TSH, which results in lower T3 and T4 release from the thyroid.
The follicular cells of the thyroid require iodides (anions of iodine) in order to synthesize T3 and T4. Iodides obtained from the diet are actively transported into follicle cells resulting in a concentration that is approximately 30 times higher than in blood. The typical diet in North America provides more iodine than required due to the addition of iodide to table salt. Inadequate iodine intake, which occurs in many developing countries, results in an inability to synthesize T3 and T4 hormones. The thyroid gland enlarges in a condition called goiter, which is caused by overproduction of TSH without the formation of thyroid hormone. Thyroglobulin is contained in a fluid called colloid, and TSH stimulation results in higher levels of colloid accumulation in the thyroid. In the absence of iodine, this is not converted to thyroid hormone, and colloid begins to accumulate more and more in the thyroid gland, leading to goiter.
Disorders can arise from both the underproduction and overproduction of thyroid hormones. Hypothyroidism, underproduction of the thyroid hormones, can cause a low metabolic rate leading to weight gain, sensitivity to cold, and reduced mental activity, among other symptoms. In children, hypothyroidism can cause cretinism, which can lead to mental retardation and growth defects. Hyperthyroidism, the overproduction of thyroid hormones, can lead to an increased metabolic rate and its effects: weight loss, excess heat production, sweating, and an increased heart rate. Graves’ disease is one example of a hyperthyroid condition.
Hormonal Control of Blood Calcium Levels
Regulation of blood calcium concentrations is important for generation of muscle contractions and nerve impulses, which are electrically stimulated. If calcium levels get too high, membrane permeability to sodium decreases and membranes become less responsive. If calcium levels get too low, membrane permeability to sodium increases and convulsions or muscle spasms can result.
Blood calcium levels are regulated by parathyroid hormone (PTH), which is produced by the parathyroid glands, as illustrated in Figure \(6\). PTH is released in response to low blood Ca2+ levels. PTH increases Ca2+ levels by targeting the skeleton, the kidneys, and the intestine. In the skeleton, PTH stimulates osteoclasts, which causes bone to be reabsorbed, releasing Ca2+ from bone into the blood. PTH also inhibits osteoblasts, reducing Ca2+ deposition in bone. In the intestines, PTH increases dietary Ca2+ absorption, and in the kidneys, PTH stimulates reabsorption of the CA2+. While PTH acts directly on the kidneys to increase Ca2+ reabsorption, its effects on the intestine are indirect. PTH triggers the formation of calcitriol, an active form of vitamin D, which acts on the intestines to increase absorption of dietary calcium. PTH release is inhibited by rising blood calcium levels.
Hyperparathyroidism results from an overproduction of parathyroid hormone. This results in excessive calcium being removed from bones and introduced into blood circulation, producing structural weakness of the bones, which can lead to deformation and fractures, plus nervous system impairment due to high blood calcium levels. Hypoparathyroidism, the underproduction of PTH, results in extremely low levels of blood calcium, which causes impaired muscle function and may result in tetany (severe sustained muscle contraction).
The hormone calcitonin, which is produced by the parafollicular or C cells of the thyroid, has the opposite effect on blood calcium levels as does PTH. Calcitonin decreases blood calcium levels by inhibiting osteoclasts, stimulating osteoblasts, and stimulating calcium excretion by the kidneys. This results in calcium being added to the bones to promote structural integrity. Calcitonin is most important in children (when it stimulates bone growth), during pregnancy (when it reduces maternal bone loss), and during prolonged starvation (because it reduces bone mass loss). In healthy nonpregnant, unstarved adults, the role of calcitonin is unclear.
Hormonal Regulation of Growth
Hormonal regulation is required for the growth and replication of most cells in the body. Growth hormone (GH), produced by the anterior portion of the pituitary gland, accelerates the rate of protein synthesis, particularly in skeletal muscle and bones. Growth hormone has direct and indirect mechanisms of action. The first direct action of GH is stimulation of triglyceride breakdown (lipolysis) and release into the blood by adipocytes. This results in a switch by most tissues from utilizing glucose as an energy source to utilizing fatty acids. This process is called a glucose-sparing effect. In another direct mechanism, GH stimulates glycogen breakdown in the liver; the glycogen is then released into the blood as glucose. Blood glucose levels increase as most tissues are utilizing fatty acids instead of glucose for their energy needs. The GH mediated increase in blood glucose levels is called a diabetogenic effect because it is similar to the high blood glucose levels seen in diabetes mellitus.
The indirect mechanism of GH action is mediated by insulin-like growth factors (IGFs) or somatomedins, which are a family of growth-promoting proteins produced by the liver, which stimulates tissue growth. IGFs stimulate the uptake of amino acids from the blood, allowing the formation of new proteins, particularly in skeletal muscle cells, cartilage cells, and other target cells, as shown in Figure \(7\). This is especially important after a meal, when glucose and amino acid concentration levels are high in the blood. GH levels are regulated by two hormones produced by the hypothalamus. GH release is stimulated by growth hormone-releasing hormone (GHRH) and is inhibited by growth hormone-inhibiting hormone (GHIH), also called somatostatin.
A balanced production of growth hormone is critical for proper development. Underproduction of GH in adults does not appear to cause any abnormalities, but in children it can result in pituitary dwarfism, in which growth is reduced. Pituitary dwarfism is characterized by symmetric body formation. In some cases, individuals are under 30 inches in height. Oversecretion of growth hormone can lead to gigantism in children, causing excessive growth. In some documented cases, individuals can reach heights of over eight feet. In adults, excessive GH can lead to acromegaly, a condition in which there is enlargement of bones in the face, hands, and feet that are still capable of growth.
Hormonal Regulation of Stress
When a threat or danger is perceived, the body responds by releasing hormones that will ready it for the “fight-or-flight” response. The effects of this response are familiar to anyone who has been in a stressful situation: increased heart rate, dry mouth, and hair standing up.
Evolution Connection: Fight-or-Flight Response
Interactions of the endocrine hormones have evolved to ensure the body’s internal environment remains stable. Stressors are stimuli that disrupt homeostasis. The sympathetic division of the vertebrate autonomic nervous system has evolved the fight-or-flight response to counter stress-induced disruptions of homeostasis. In the initial alarm phase, the sympathetic nervous system stimulates an increase in energy levels through increased blood glucose levels. This prepares the body for physical activity that may be required to respond to stress: to either fight for survival or to flee from danger.
However, some stresses, such as illness or injury, can last for a long time. Glycogen reserves, which provide energy in the short-term response to stress, are exhausted after several hours and cannot meet long-term energy needs. If glycogen reserves were the only energy source available, neural functioning could not be maintained once the reserves became depleted due to the nervous system’s high requirement for glucose. In this situation, the body has evolved a response to counter long-term stress through the actions of the glucocorticoids, which ensure that long-term energy requirements can be met. The glucocorticoids mobilize lipid and protein reserves, stimulate gluconeogenesis, conserve glucose for use by neural tissue, and stimulate the conservation of salts and water. The mechanisms to maintain homeostasis that are described here are those observed in the human body. However, the fight-or-flight response exists in some form in all vertebrates.
The sympathetic nervous system regulates the stress response via the hypothalamus. Stressful stimuli cause the hypothalamus to signal the adrenal medulla (which mediates short-term stress responses) via nerve impulses, and the adrenal cortex, which mediates long-term stress responses, via the hormone adrenocorticotropic hormone (ACTH), which is produced by the anterior pituitary.
Short-term Stress Response
When presented with a stressful situation, the body responds by calling for the release of hormones that provide a burst of energy. The hormones epinephrine (also known as adrenaline) and norepinephrine (also known as noradrenaline) are released by the adrenal medulla. How do these hormones provide a burst of energy? Epinephrine and norepinephrine increase blood glucose levels by stimulating the liver and skeletal muscles to break down glycogen and by stimulating glucose release by liver cells. Additionally, these hormones increase oxygen availability to cells by increasing the heart rate and dilating the bronchioles. The hormones also prioritize body function by increasing blood supply to essential organs such as the heart, brain, and skeletal muscles, while restricting blood flow to organs not in immediate need, such as the skin, digestive system, and kidneys. Epinephrine and norepinephrine are collectively called catecholamines.
Link to Learning
Watch this Discovery Channel animation describing the flight-or-flight response.
Long-term Stress Response
Long-term stress response differs from short-term stress response. The body cannot sustain the bursts of energy mediated by epinephrine and norepinephrine for long times. Instead, other hormones come into play. In a long-term stress response, the hypothalamus triggers the release of ACTH from the anterior pituitary gland. The adrenal cortex is stimulated by ACTH to release steroid hormones called corticosteroids. Corticosteroids turn on transcription of certain genes in the nuclei of target cells. They change enzyme concentrations in the cytoplasm and affect cellular metabolism. There are two main corticosteroids: glucocorticoids such as cortisol, and mineralocorticoids such as aldosterone. These hormones target the breakdown of fat into fatty acids in the adipose tissue. The fatty acids are released into the bloodstream for other tissues to use for ATP production. The glucocorticoids primarily affect glucose metabolism by stimulating glucose synthesis. Glucocorticoids also have anti-inflammatory properties through inhibition of the immune system. For example, cortisone is used as an anti-inflammatory medication; however, it cannot be used long term as it increases susceptibility to disease due to its immune-suppressing effects.
Mineralocorticoids function to regulate ion and water balance of the body. The hormone aldosterone stimulates the reabsorption of water and sodium ions in the kidney, which results in increased blood pressure and volume.
Hypersecretion of glucocorticoids can cause a condition known as Cushing’s disease, characterized by a shifting of fat storage areas of the body. This can cause the accumulation of adipose tissue in the face and neck, and excessive glucose in the blood. Hyposecretion of the corticosteroids can cause Addison’s disease, which may result in bronzing of the skin, hypoglycemia, and low electrolyte levels in the blood.
Summary
Water levels in the body are controlled by antidiuretic hormone (ADH), which is produced in the hypothalamus and triggers the reabsorption of water by the kidneys. Underproduction of ADH can cause diabetes insipidus. Aldosterone, a hormone produced by the adrenal cortex of the kidneys, enhances Na+ reabsorption from the extracellular fluids and subsequent water reabsorption by diffusion. The renin-angiotensin-aldosterone system is one way that aldosterone release is controlled.
The reproductive system is controlled by the gonadotropins follicle-stimulating hormone (FSH) and luteinizing hormone (LH), which are produced by the pituitary gland. Gonadotropin release is controlled by the hypothalamic hormone gonadotropin-releasing hormone (GnRH). FSH stimulates the maturation of sperm cells in males and is inhibited by the hormone inhibin, while LH stimulates the production of the androgen testosterone. FSH stimulates egg maturation in females, while LH stimulates the production of estrogens and progesterone. Estrogens are a group of steroid hormones produced by the ovaries that trigger the development of secondary sex characteristics in females as well as control the maturation of the ova. In females, the pituitary also produces prolactin, which stimulates milk production after childbirth, and oxytocin, which stimulates uterine contraction during childbirth and milk let-down during suckling.
Insulin is produced by the pancreas in response to rising blood glucose levels and allows cells to utilize blood glucose and store excess glucose for later use. Diabetes mellitus is caused by reduced insulin activity and causes high blood glucose levels, or hyperglycemia. Glucagon is released by the pancreas in response to low blood glucose levels and stimulates the breakdown of glycogen into glucose, which can be used by the body. The body’s basal metabolic rate is controlled by the thyroid hormones thyroxine (T4) and triiodothyronine (T3). The anterior pituitary produces thyroid stimulating hormone (TSH), which controls the release of T3 and T4 from the thyroid gland. Iodine is necessary in the production of thyroid hormone, and the lack of iodine can lead to a condition called goiter.
Parathyroid hormone (PTH) is produced by the parathyroid glands in response to low blood Ca2+ levels. The parafollicular cells of the thyroid produce calcitonin, which reduces blood Ca2+ levels. Growth hormone (GH) is produced by the anterior pituitary and controls the growth rate of muscle and bone. GH action is indirectly mediated by insulin-like growth factors (IGFs). Short-term stress causes the hypothalamus to trigger the adrenal medulla to release epinephrine and norepinephrine, which trigger the fight or flight response. Long-term stress causes the hypothalamus to trigger the anterior pituitary to release adrenocorticotropic hormone (ACTH), which causes the release of corticosteroids, glucocorticoids, and mineralocorticoids, from the adrenal cortex.
Glossary
acromegaly
condition caused by overproduction of GH in adults
Addison’s disease
disorder caused by the hyposecretion of corticosteroids
adrenocorticotropic hormone (ACTH)
hormone released by the anterior pituitary, which stimulates the adrenal cortex to release corticosteroids during the long-term stress response
aldosterone
steroid hormone produced by the adrenal cortex that stimulates the reabsorption of Na+ from extracellular fluids and secretion of K+.
androgen
male sex hormone such as testosterone
antidiuretic hormone (ADH)
hormone produced by the hypothalamus and released by the posterior pituitary that increases water reabsorption by the kidneys
calcitonin
hormone produced by the parafollicular cells of the thyroid gland that functions to lower blood Ca2+ levels and promote bone growth
corticosteroid
hormone released by the adrenal cortex in response to long-term stress
cortisol
glucocorticoid produced in response to stress
Cushing’s disease
disorder caused by the hypersecretion of glucocorticoids
diabetes insipidus
disorder caused by underproduction of ADH
diabetes mellitus
disorder caused by low levels of insulin activity
diabetogenic effect
effect of GH that causes blood glucose levels to rise similar to diabetes mellitus
epinephrine
hormone released by the adrenal medulla in response to a short term stress
estrogens
- a group of steroid hormones, including estradiol and several others, that are produced by the ovaries and elicit secondary sex characteristics in females as well as control the maturation of the ova
follicle-stimulating hormone (FSH)
hormone produced by the anterior pituitary that stimulates gamete production
gigantism
condition caused by overproduction of GH in children
glucagon
hormone produced by the alpha cells of the pancreas in response to low blood sugar; functions to raise blood sugar levels
glucocorticoid
corticosteroid that affects glucose metabolism
gluconeogenesis
synthesis of glucose from amino acids
glucose-sparing effect
effect of GH that causes tissues to use fatty acids instead of glucose as an energy source
glycogenolysis
breakdown of glycogen into glucose
goiter
enlargement of the thyroid gland caused by insufficient dietary iodine levels
gonadotropin
hormone that regulates the gonads, including FSH and LH
growth hormone (GH)
hormone produced by the anterior pituitary that promotes protein synthesis and body growth
growth hormone-inhibiting hormone (GHIH)
hormone produced by the hypothalamus that inhibits growth hormone production, also called somatostatin
growth hormone-releasing hormone (GHRH)
hormone released by the hypothalamus that triggers the release of GH
hyperglycemia
high blood sugar level
hyperthyroidism
overactivity of the thyroid gland
hypoglycemia
low blood sugar level
hypothyroidism
underactivity of the thyroid gland
insulin
hormone produced by the beta cells of the pancreas in response to high blood glucose levels; functions to lower blood glucose levels
insulin-like growth factor (IGF)
growth-promoting protein produced by the liver
mineralocorticoid
corticosteroid that affects ion and water balance
norepinephrine
hormone released by the adrenal medulla in response to a short-term stress hormone production by the gonads
osmoreceptor
receptor in the hypothalamus that monitors the concentration of electrolytes in the blood
oxytocin
hormone released by the posterior pituitary to stimulate uterine contractions during childbirth and milk let-down in the mammary glands
parathyroid hormone (PTH)
hormone produced by the parathyroid glands in response to low blood Ca2+ levels; functions to raise blood Ca2+ levels
pituitary dwarfism
condition caused by underproduction of GH in children
prolactin (PRL)
hormone produced by the anterior pituitary that stimulates milk production
prolactin-inhibiting hormone
hormone produced by the hypothalamus that inhibits the release of prolactin
prolactin-releasing hormone
hormone produced by the hypothalamus that stimulates the release of prolactin
renin
enzyme produced by the juxtaglomerular apparatus of the kidneys that reacts with angiotensinogen to cause the release of aldosterone
thyroglobulin
glycoprotein found in the thyroid that is converted into thyroid hormone
thyroid-stimulating hormone (TSH)
hormone produced by the anterior pituitary that controls the release of T3 and T4 from the thyroid gland
thyroxine (tetraiodothyronine, T4)
thyroid hormone that controls the basal metabolic rate
triiodothyronine (T3)
thyroid hormone that controls the basal metabolic rate | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.03%3A_The_Pituitary_and_Hypothalamus-_The_Body%27s_Control_Centers/44.3.01%3A_Regulation_of_Body_Processes.txt |
Skills to Develop
• Describe the role of different glands in the endocrine system
• Explain how the different glands work together to maintain homeostasis
Both the endocrine and nervous systems use chemical signals to communicate and regulate the body's physiology. The endocrine system releases hormones that act on target cells to regulate development, growth, energy metabolism, reproduction, and many behaviors. The nervous system releases neurotransmitters or neurohormones that regulate neurons, muscle cells, and endocrine cells. Because the neurons can regulate the release of hormones, the nervous and endocrine systems work in a coordinated manner to regulate the body's physiology.
Hypothalamic-Pituitary Axis
The hypothalamus in vertebrates integrates the endocrine and nervous systems. The hypothalamus is an endocrine organ located in the diencephalon of the brain. It receives input from the body and other brain areas and initiates endocrine responses to environmental changes. The hypothalamus acts as an endocrine organ, synthesizing hormones and transporting them along axons to the posterior pituitary gland. It synthesizes and secretes regulatory hormones that control the endocrine cells in the anterior pituitary gland. The hypothalamus contains autonomic centers that control endocrine cells in the adrenal medulla via neuronal control.
The pituitary gland, sometimes called the hypophysis or “master gland” is located at the base of the brain in the sella turcica, a groove of the sphenoid bone of the skull, illustrated in Figure \(1\). It is attached to the hypothalamus via a stalk called the pituitary stalk (or infundibulum). The anterior portion of the pituitary gland is regulated by releasing or release-inhibiting hormones produced by the hypothalamus, and the posterior pituitary receives signals via neurosecretory cells to release hormones produced by the hypothalamus. The pituitary has two distinct regions—the anterior pituitary and the posterior pituitary—which between them secrete nine different peptide or protein hormones. The posterior lobe of the pituitary gland contains axons of the hypothalamic neurons.
Anterior Pituitary
The anterior pituitary gland, or adenohypophysis, is surrounded by a capillary network that extends from the hypothalamus, down along the infundibulum, and to the anterior pituitary. This capillary network is a part of the hypophyseal portal system that carries substances from the hypothalamus to the anterior pituitary and hormones from the anterior pituitary into the circulatory system. A portal system carries blood from one capillary network to another; therefore, the hypophyseal portal system allows hormones produced by the hypothalamus to be carried directly to the anterior pituitary without first entering the circulatory system.
The anterior pituitary produces seven hormones: growth hormone (GH), prolactin (PRL), thyroid-stimulating hormone (TSH), melanin-stimulating hormone (MSH), adrenocorticotropic hormone (ACTH), follicle-stimulating hormone (FSH), and luteinizing hormone (LH). Anterior pituitary hormones are sometimes referred to as tropic hormones, because they control the functioning of other organs. While these hormones are produced by the anterior pituitary, their production is controlled by regulatory hormones produced by the hypothalamus. These regulatory hormones can be releasing hormones or inhibiting hormones, causing more or less of the anterior pituitary hormones to be secreted. These travel from the hypothalamus through the hypophyseal portal system to the anterior pituitary where they exert their effect. Negative feedback then regulates how much of these regulatory hormones are released and how much anterior pituitary hormone is secreted.
Posterior Pituitary
The posterior pituitary is significantly different in structure from the anterior pituitary. It is a part of the brain, extending down from the hypothalamus, and contains mostly nerve fibers and neuroglial cells, which support axons that extend from the hypothalamus to the posterior pituitary. The posterior pituitary and the infundibulum together are referred to as the neurohypophysis.
The hormones antidiuretic hormone (ADH), also known as vasopressin, and oxytocin are produced by neurons in the hypothalamus and transported within these axons along the infundibulum to the posterior pituitary. They are released into the circulatory system via neural signaling from the hypothalamus. These hormones are considered to be posterior pituitary hormones, even though they are produced by the hypothalamus, because that is where they are released into the circulatory system. The posterior pituitary itself does not produce hormones, but instead stores hormones produced by the hypothalamus and releases them into the blood stream.
Thyroid Gland
The thyroid gland is located in the neck, just below the larynx and in front of the trachea, as shown in Figure \(2\). It is a butterfly-shaped gland with two lobes that are connected by the isthmus. It has a dark red color due to its extensive vascular system. When the thyroid swells due to dysfunction, it can be felt under the skin of the neck.
The thyroid gland is made up of many spherical thyroid follicles, which are lined with a simple cuboidal epithelium. These follicles contain a viscous fluid, called colloid, which stores the glycoprotein thyroglobulin, the precursor to the thyroid hormones. The follicles produce hormones that can be stored in the colloid or released into the surrounding capillary network for transport to the rest of the body via the circulatory system.
Thyroid follicle cells synthesize the hormone thyroxine, which is also known as T4 because it contains four atoms of iodine, and triiodothyronine, also known as T3 because it contains three atoms of iodine. Follicle cells are stimulated to release stored T3 and T4 by thyroid stimulating hormone (TSH), which is produced by the anterior pituitary. These thyroid hormones increase the rates of mitochondrial ATP production.
A third hormone, calcitonin, is produced by parafollicular cells of the thyroid either releasing hormones or inhibiting hormones. Calcitonin release is not controlled by TSH, but instead is released when calcium ion concentrations in the blood rise. Calcitonin functions to help regulate calcium concentrations in body fluids. It acts in the bones to inhibit osteoclast activity and in the kidneys to stimulate excretion of calcium. The combination of these two events lowers body fluid levels of calcium.
Parathyroid Glands
Most people have four parathyroid glands; however, the number can vary from two to six. These glands are located on the posterior surface of the thyroid gland, as shown in Figure \(3\). Normally, there is a superior gland and an inferior gland associated with each of the thyroid’s two lobes. Each parathyroid gland is covered by connective tissue and contains many secretory cells that are associated with a capillary network.
The parathyroid glands produce parathyroid hormone (PTH). PTH increases blood calcium concentrations when calcium ion levels fall below normal. PTH (1) enhances reabsorption of Ca2+ by the kidneys, (2) stimulates osteoclast activity and inhibits osteoblast activity, and (3) it stimulates synthesis and secretion of calcitriol by the kidneys, which enhances Ca2+ absorption by the digestive system. PTH is produced by chief cells of the parathyroid. PTH and calcitonin work in opposition to one another to maintain homeostatic Ca2+ levels in body fluids. Another type of cells, oxyphil cells, exist in the parathyroid but their function is not known. These hormones encourage bone growth, muscle mass, and blood cell formation in children and women.
Adrenal Glands
The adrenal glands are associated with the kidneys; one gland is located on top of each kidney as illustrated in Figure \(4\). The adrenal glands consist of an outer adrenal cortex and an inner adrenal medulla. These regions secrete different hormones.
Adrenal Cortex
The adrenal cortex is made up of layers of epithelial cells and associated capillary networks. These layers form three distinct regions: an outer zona glomerulosa that produces mineralocorticoids, a middle zona fasciculata that produces glucocorticoids, and an inner zona reticularis that produces androgens.
The main mineralocorticoid is aldosterone, which regulates the concentration of Na+ ions in urine, sweat, pancreas, and saliva. Aldosterone release from the adrenal cortex is stimulated by a decrease in blood concentrations of sodium ions, blood volume, or blood pressure, or by an increase in blood potassium levels.
The three main glucocorticoids are cortisol, corticosterone, and cortisone. The glucocorticoids stimulate the synthesis of glucose and gluconeogenesis (converting a non-carbohydrate to glucose) by liver cells and they promote the release of fatty acids from adipose tissue. These hormones increase blood glucose levels to maintain levels within a normal range between meals. These hormones are secreted in response to ACTH and levels are regulated by negative feedback.
Androgens are sex hormones that promote masculinity. They are produced in small amounts by the adrenal cortex in both males and females. They do not affect sexual characteristics and may supplement sex hormones released from the gonads.
Adrenal Medulla
The adrenal medulla contains large, irregularly shaped cells that are closely associated with blood vessels. These cells are innervated by preganglionic autonomic nerve fibers from the central nervous system.
The adrenal medulla contains two types of secretory cells: one that produces epinephrine (adrenaline) and another that produces norepinephrine (noradrenaline). Epinephrine is the primary adrenal medulla hormone accounting for 75 to 80 percent of its secretions. Epinephrine and norepinephrine increase heart rate, breathing rate, cardiac muscle contractions, blood pressure, and blood glucose levels. They also accelerate the breakdown of glucose in skeletal muscles and stored fats in adipose tissue.
The release of epinephrine and norepinephrine is stimulated by neural impulses from the sympathetic nervous system. Secretion of these hormones is stimulated by acetylcholine release from preganglionic sympathetic fibers innervating the adrenal medulla. These neural impulses originate from the hypothalamus in response to stress to prepare the body for the fight-or-flight response.
Pancreas
The pancreas, illustrated in Figure \(5\), is an elongated organ that is located between the stomach and the proximal portion of the small intestine. It contains both exocrine cells that excrete digestive enzymes and endocrine cells that release hormones. It is sometimes referred to as a heterocrine gland because it has both endocrine and exocrine functions.
The endocrine cells of the pancreas form clusters called pancreatic islets or the islets of Langerhans, as visible in the micrograph shown in Figure \(6\). The pancreatic islets contain two primary cell types: alpha cells, which produce the hormone glucagon, and beta cells, which produce the hormone insulin. These hormones regulate blood glucose levels. As blood glucose levels decline, alpha cells release glucagon to raise the blood glucose levels by increasing rates of glycogen breakdown and glucose release by the liver. When blood glucose levels rise, such as after a meal, beta cells release insulin to lower blood glucose levels by increasing the rate of glucose uptake in most body cells, and by increasing glycogen synthesis in skeletal muscles and the liver. Together, glucagon and insulin regulate blood glucose levels.
Pineal Gland
The pineal gland produces melatonin. The rate of melatonin production is affected by the photoperiod. Collaterals from the visual pathways innervate the pineal gland. During the day photoperiod, little melatonin is produced; however, melatonin production increases during the dark photoperiod (night). In some mammals, melatonin has an inhibitory affect on reproductive functions by decreasing production and maturation of sperm, oocytes, and reproductive organs. Melatonin is an effective antioxidant, protecting the CNS from free radicals such as nitric oxide and hydrogen peroxide. Lastly, melatonin is involved in biological rhythms, particularly circadian rhythms such as the sleep-wake cycle and eating habits.
Gonads
The gonads—the male testes and female ovaries—produce steroid hormones. The testes produce androgens, testosterone being the most prominent, which allow for the development of secondary sex characteristics and the production of sperm cells. The ovaries produce estradiol and progesterone, which cause secondary sex characteristics and prepare the body for childbirth.
Table \(1\): Endocrine Glands and their Associated Hormones
Endocrine Gland Associated Hormones Effect
Hypothalamus releasing and inhibiting hormones regulate hormone release from pituitary gland; produce oxytocin; produce uterine contractions and milk secretion in females
antidiuretic hormone (ADH) water reabsorption from kidneys; vasoconstriction to increase blood pressure
Pituitary (Anterior) growth hormone (GH) promotes growth of body tissues, protein synthesis; metabolic functions
prolactin (PRL) promotes milk production
thyroid stimulating hormone (TSH) stimulates thyroid hormone release
adrenocorticotropic hormone (ACTH) stimulates hormone release by adrenal cortex, glucocorticoids
follicle-stimulating hormone (FSH) stimulates gamete production (both ova and sperm); secretion of estradiol
luteinizing hormone (LH) stimulates androgen production by gonads; ovulation, secretion of progesterone
melanocyte-stimulating hormone (MSH) stimulates melanocytes of the skin increasing melanin pigment production.
Pituitary (Posterior) antidiuretic hormone (ADH) stimulates water reabsorption by kidneys
oxytocin stimulates uterine contractions during childbirth; milk ejection; stimulates ductus deferens and prostate gland contraction during emission
Thyroid thyroxine, triiodothyronine stimulate and maintain metabolism; growth and development
calcitonin reduces blood Ca2+ levels
Parathyroid parathyroid hormone (PTH) increases blood Ca2+ levels
Adrenal (Cortex) aldosterone increases blood Na+ levels; increase K+ secretion
cortisol, corticosterone, cortisone increase blood glucose levels; anti-inflammatory effects
Adrenal (Medulla) epinephrine, norepinephrine stimulate fight-or-flight response; increase blood gluclose levels; increase metabolic activities
Pancreas insulin reduces blood glucose levels
glucagon increases blood glucose levels
Pineal gland melatonin regulates some biological rhythms and protects CNS from free radicals
Testes androgens regulate, promote, increase or maintain sperm production; male secondary sexual characteristics
Ovaries estrogen promotes uterine lining growth; female secondary sexual characteristics
progestins promote and maintain uterine lining growth
Organs with Secondary Endocrine Functions
There are several organs whose primary functions are non-endocrine but that also possess endocrine functions. These include the heart, kidneys, intestines, thymus, gonads, and adipose tissue.
The heart possesses endocrine cells in the walls of the atria that are specialized cardiac muscle cells. These cells release the hormone atrial natriuretic peptide (ANP) in response to increased blood volume. High blood volume causes the cells to be stretched, resulting in hormone release. ANP acts on the kidneys to reduce the reabsorption of Na+, causing Na+ and water to be excreted in the urine. ANP also reduces the amounts of renin released by the kidneys and aldosterone released by the adrenal cortex, further preventing the retention of water. In this way, ANP causes a reduction in blood volume and blood pressure, and reduces the concentration of Na+ in the blood.
The gastrointestinal tract produces several hormones that aid in digestion. The endocrine cells are located in the mucosa of the GI tract throughout the stomach and small intestine. Some of the hormones produced include gastrin, secretin, and cholecystokinin, which are secreted in the presence of food, and some of which act on other organs such as the pancreas, gallbladder, and liver. They trigger the release of gastric juices, which help to break down and digest food in the GI tract.
While the adrenal glands associated with the kidneys are major endocrine glands, the kidneys themselves also possess endocrine function. Renin is released in response to decreased blood volume or pressure and is part of the renin-angiotensin-aldosterone system that leads to the release of aldosterone. Aldosterone then causes the retention of Na+ and water, raising blood volume. The kidneys also release calcitriol, which aids in the absorption of Ca2+ and phosphate ions. Erythropoietin (EPO) is a protein hormone that triggers the formation of red blood cells in the bone marrow. EPO is released in response to low oxygen levels. Because red blood cells are oxygen carriers, increased production results in greater oxygen delivery throughout the body. EPO has been used by athletes to improve performance, as greater oxygen delivery to muscle cells allows for greater endurance. Because red blood cells increase the viscosity of blood, artificially high levels of EPO can cause severe health risks.
The thymus is found behind the sternum; it is most prominent in infants, becoming smaller in size through adulthood. The thymus produces hormones referred to as thymosins, which contribute to the development of the immune response.
Adipose tissue is a connective tissue found throughout the body. It produces the hormone leptin in response to food intake. Leptin increases the activity of anorexigenic neurons and decreases that of orexigenic neurons, producing a feeling of satiety after eating, thus affecting appetite and reducing the urge for further eating. Leptin is also associated with reproduction. It must be present for GnRH and gonadotropin synthesis to occur. Extremely thin females may enter puberty late; however, if adipose levels increase, more leptin will be produced, improving fertility.
Summary
The pituitary gland is located at the base of the brain and is attached to the hypothalamus by the infundibulum. The anterior pituitary receives products from the hypothalamus by the hypophyseal portal system and produces six hormones. The posterior pituitary is an extension of the brain and releases hormones (antidiuretic hormone and oxytocin) produced by the hypothalamus.
The thyroid gland is located in the neck and is composed of two lobes connected by the isthmus. The thyroid is made up of follicle cells that produce the hormones thyroxine and triiodothyronine. Parafollicular cells of the thyroid produce calcitonin. The parathyroid glands lie on the posterior surface of the thyroid gland and produce parathyroid hormone.
The adrenal glands are located on top of the kidneys and consist of the renal cortex and renal medulla. The adrenal cortex is the outer part of the adrenal gland and produces the corticosteroids, glucocorticoids, and mineralocorticoids. The adrenal medulla is the inner part of the adrenal gland and produces the catecholamines epinephrine and norepinephrine.
The pancreas lies in the abdomen between the stomach and the small intestine. Clusters of endocrine cells in the pancreas form the islets of Langerhans, which are composed of alpha cells that release glucagon and beta cells that release insulin.
Some organs possess endocrine activity as a secondary function but have another primary function. The heart produces the hormone atrial natriuretic peptide, which functions to reduce blood volume, pressure, and Na+ concentration. The gastrointestinal tract produces various hormones that aid in digestion. The kidneys produce renin, calcitriol, and erythropoietin. Adipose tissue produces leptin, which promotes satiety signals in the brain.
Glossary
adrenal cortex
outer portion of adrenal glands that produces corticosteroids
adrenal gland
endocrine glands associated with the kidneys
adrenal medulla
inner portion of adrenal glands that produces epinephrine and norepinephrine
alpha cell
endocrine cell of the pancreatic islets that produces the hormone glucagon
anterior pituitary
portion of the pituitary gland that produces six hormones; also called adenohypophysis
atrial natriuretic peptide (ANP)
hormone produced by the heart to reduce blood volume, pressure, and Na+ concentration
beta cell
endocrine cell of the pancreatic islets that produces the hormone insulin
colloid
fluid inside the thyroid gland that contains the glycoprotein thyroglobulin
endocrine gland
gland that secretes hormones into the surrounding interstitial fluid, which then diffuse into blood and are carried to various organs and tissues within the body
erythropoietin (EPO)
hormone produced by the kidneys to stimulate red blood cell production in the bone marrow
hypophyseal portal system
system of blood vessels that carries hormones from the hypothalamus to the anterior pituitary
islets of Langerhans (pancreatic islets)
endocrine cells of the pancreas
isthmus
tissue mass that connects the two lobes of the thyroid gland
leptin
hormone produced by adipose tissue that promotes feelings of satiety and reduces hunger
pancreas
organ located between the stomach and the small intestine that contains exocrine and endocrine cells
parafollicular cell
thyroid cell that produces the hormone calcitonin
parathyroid gland
gland located on the surface of the thyroid that produces parathyroid hormone
pituitary gland
endocrine gland located at the base of the brain composed of an anterior and posterior region; also called hypophysis
pituitary stalk
(also, infundibulum) stalk that connects the pituitary gland to the hypothalamus
posterior pituitary
extension of the brain that releases hormones produced by the hypothalamus; along with the infundibulum, it is also referred to as the neurohypophysis
thymus
gland located behind the sternum that produces thymosin hormones that contribute to the development of the immune system
thyroid gland
endocrine gland located in the neck that produces thyroid hormones thyroxine and triiodothyronine | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.03%3A_The_Pituitary_and_Hypothalamus-_The_Body%27s_Control_Centers/44.3.02%3A_Endocrine_Glands.txt |
Skills to Develop
• Explain how hormones regulate the excretory system
• Discuss the role of hormones in the reproductive system
• Describe how hormones regulate metabolism
• Explain the role of hormones in different diseases
Hormones have a wide range of effects and modulate many different body processes. The key regulatory processes that will be examined here are those affecting the excretory system, the reproductive system, metabolism, blood calcium concentrations, growth, and the stress response.
Hormonal Regulation of the Excretory System
Maintaining a proper water balance in the body is important to avoid dehydration or over-hydration (hyponatremia). The water concentration of the body is monitored by osmoreceptors in the hypothalamus, which detect the concentration of electrolytes in the extracellular fluid. The concentration of electrolytes in the blood rises when there is water loss caused by excessive perspiration, inadequate water intake, or low blood volume due to blood loss. An increase in blood electrolyte levels results in a neuronal signal being sent from the osmoreceptors in hypothalamic nuclei. The pituitary gland has two components: anterior and posterior. The anterior pituitary is composed of glandular cells that secrete protein hormones. The posterior pituitary is an extension of the hypothalamus. It is composed largely of neurons that are continuous with the hypothalamus.
The hypothalamus produces a polypeptide hormone known as antidiuretic hormone (ADH), which is transported to and released from the posterior pituitary gland. The principal action of ADH is to regulate the amount of water excreted by the kidneys. As ADH (which is also known as vasopressin) causes direct water reabsorption from the kidney tubules, salts and wastes are concentrated in what will eventually be excreted as urine. The hypothalamus controls the mechanisms of ADH secretion, either by regulating blood volume or the concentration of water in the blood. Dehydration or physiological stress can cause an increase of osmolarity above 300 mOsm/L, which in turn, raises ADH secretion and water will be retained, causing an increase in blood pressure. ADH travels in the bloodstream to the kidneys. Once at the kidneys, ADH changes the kidneys to become more permeable to water by temporarily inserting water channels, aquaporins, into the kidney tubules. Water moves out of the kidney tubules through the aquaporins, reducing urine volume. The water is reabsorbed into the capillaries lowering blood osmolarity back toward normal. As blood osmolarity decreases, a negative feedback mechanism reduces osmoreceptor activity in the hypothalamus, and ADH secretion is reduced. ADH release can be reduced by certain substances, including alcohol, which can cause increased urine production and dehydration.
Chronic underproduction of ADH or a mutation in the ADH receptor results in diabetes insipidus. If the posterior pituitary does not release enough ADH, water cannot be retained by the kidneys and is lost as urine. This causes increased thirst, but water taken in is lost again and must be continually consumed. If the condition is not severe, dehydration may not occur, but severe cases can lead to electrolyte imbalances due to dehydration.
Another hormone responsible for maintaining electrolyte concentrations in extracellular fluids is aldosterone, a steroid hormone that is produced by the adrenal cortex. In contrast to ADH, which promotes the reabsorption of water to maintain proper water balance, aldosterone maintains proper water balance by enhancing Na+ reabsorption and K+ secretion from extracellular fluid of the cells in kidney tubules. Because it is produced in the cortex of the adrenal gland and affects the concentrations of minerals Na+ and K+, aldosterone is referred to as a mineralocorticoid, a corticosteroid that affects ion and water balance. Aldosterone release is stimulated by a decrease in blood sodium levels, blood volume, or blood pressure, or an increase in blood potassium levels. It also prevents the loss of Na+ from sweat, saliva, and gastric juice. The reabsorption of Na+ also results in the osmotic reabsorption of water, which alters blood volume and blood pressure.
Aldosterone production can be stimulated by low blood pressure, which triggers a sequence of chemical release, as illustrated in Figure \(1\). When blood pressure drops, the renin-angiotensin-aldosterone system (RAAS) is activated. Cells in the juxtaglomerular apparatus, which regulates the functions of the nephrons of the kidney, detect this and release renin. Renin, an enzyme, circulates in the blood and reacts with a plasma protein produced by the liver called angiotensinogen. When angiotensinogen is cleaved by renin, it produces angiotensin I, which is then converted into angiotensin II in the lungs. Angiotensin II functions as a hormone and then causes the release of the hormone aldosterone by the adrenal cortex, resulting in increased Na+ reabsorption, water retention, and an increase in blood pressure. Angiotensin II in addition to being a potent vasoconstrictor also causes an increase in ADH and increased thirst, both of which help to raise blood pressure.
Hormonal Regulation of the Reproductive System
Regulation of the reproductive system is a process that requires the action of hormones from the pituitary gland, the adrenal cortex, and the gonads. During puberty in both males and females, the hypothalamus produces gonadotropin-releasing hormone (GnRH), which stimulates the production and release of follicle-stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary gland. These hormones regulate the gonads (testes in males and ovaries in females) and therefore are called gonadotropins. In both males and females, FSH stimulates gamete production and LH stimulates production of hormones by the gonads. An increase in gonad hormone levels inhibits GnRH production through a negative feedback loop.
Regulation of the Male Reproductive System
In males, FSH stimulates the maturation of sperm cells. FSH production is inhibited by the hormone inhibin, which is released by the testes. LH stimulates production of the sex hormones (androgens) by the interstitial cells of the testes and therefore is also called interstitial cell-stimulating hormone. The most widely known androgen in males is testosterone. Testosterone promotes the production of sperm and masculine characteristics. The adrenal cortex also produces small amounts of testosterone precursor, although the role of this additional hormone production is not fully understood.
Everyday Connection: The Dangers of Synthetic Hormones
Some athletes attempt to boost their performance by using artificial hormones that enhance muscle performance. Anabolic steroids, a form of the male sex hormone testosterone, are one of the most widely known performance-enhancing drugs. Steroids are used to help build muscle mass. Other hormones that are used to enhance athletic performance include erythropoietin, which triggers the production of red blood cells, and human growth hormone, which can help in building muscle mass. Most performance enhancing drugs are illegal for non-medical purposes. They are also banned by national and international governing bodies including the International Olympic Committee, the U.S. Olympic Committee, the National Collegiate Athletic Association, the Major League Baseball, and the National Football League.
The side effects of synthetic hormones are often significant and non-reversible, and in some cases, fatal. Androgens produce several complications such as liver dysfunctions and liver tumors, prostate gland enlargement, difficulty urinating, premature closure of epiphyseal cartilages, testicular atrophy, infertility, and immune system depression. The physiological strain caused by these substances is often greater than what the body can handle, leading to unpredictable and dangerous effects and linking their use to heart attacks, strokes, and impaired cardiac function.
Regulation of the Female Reproductive System
In females, FSH stimulates development of egg cells, called ova, which develop in structures called follicles. Follicle cells produce the hormone inhibin, which inhibits FSH production. LH also plays a role in the development of ova, induction of ovulation, and stimulation of estradiol and progesterone production by the ovaries, as illustrated in Figure \(3\). Estradiol and progesterone are steroid hormones that prepare the body for pregnancy. Estradiol produces secondary sex characteristics in females, while both estradiol and progesterone regulate the menstrual cycle.
In addition to producing FSH and LH, the anterior portion of the pituitary gland also produces the hormone prolactin (PRL) in females. Prolactin stimulates the production of milk by the mammary glands following childbirth. Prolactin levels are regulated by the hypothalamic hormones prolactin-releasing hormone (PRH) and prolactin-inhibiting hormone (PIH), which is now known to be dopamine. PRH stimulates the release of prolactin and PIH inhibits it.
The posterior pituitary releases the hormone oxytocin, which stimulates uterine contractions during childbirth. The uterine smooth muscles are not very sensitive to oxytocin until late in pregnancy when the number of oxytocin receptors in the uterus peaks. Stretching of tissues in the uterus and cervix stimulates oxytocin release during childbirth. Contractions increase in intensity as blood levels of oxytocin rise via a positive feedback mechanism until the birth is complete. Oxytocin also stimulates the contraction of myoepithelial cells around the milk-producing mammary glands. As these cells contract, milk is forced from the secretory alveoli into milk ducts and is ejected from the breasts in milk ejection (“let-down”) reflex. Oxytocin release is stimulated by the suckling of an infant, which triggers the synthesis of oxytocin in the hypothalamus and its release into circulation at the posterior pituitary.
Hormonal Regulation of Metabolism
Blood glucose levels vary widely over the course of a day as periods of food consumption alternate with periods of fasting. Insulin and glucagon are the two hormones primarily responsible for maintaining homeostasis of blood glucose levels. Additional regulation is mediated by the thyroid hormones.
Regulation of Blood Glucose Levels by Insulin and Glucagon
Cells of the body require nutrients in order to function, and these nutrients are obtained through feeding. In order to manage nutrient intake, storing excess intake and utilizing reserves when necessary, the body uses hormones to moderate energy stores. Insulin is produced by the beta cells of the pancreas, which are stimulated to release insulin as blood glucose levels rise (for example, after a meal is consumed). Insulin lowers blood glucose levels by enhancing the rate of glucose uptake and utilization by target cells, which use glucose for ATP production. It also stimulates the liver to convert glucose to glycogen, which is then stored by cells for later use. Insulin also increases glucose transport into certain cells, such as muscle cells and the liver. This results from an insulin-mediated increase in the number of glucose transporter proteins in cell membranes, which remove glucose from circulation by facilitated diffusion. As insulin binds to its target cell via insulin receptors and signal transduction, it triggers the cell to incorporate glucose transport proteins into its membrane. This allows glucose to enter the cell, where it can be used as an energy source. However, this does not occur in all cells: some cells, including those in the kidneys and brain, can access glucose without the use of insulin. Insulin also stimulates the conversion of glucose to fat in adipocytes and the synthesis of proteins. These actions mediated by insulin cause blood glucose concentrations to fall, called a hypoglycemic “low sugar” effect, which inhibits further insulin release from beta cells through a negative feedback loop.
Link to Learning
This animation describes the role of insulin and the pancreas in diabetes.
Impaired insulin function can lead to a condition called diabetes mellitus, the main symptoms of which are illustrated in Figure \(4\). This can be caused by low levels of insulin production by the beta cells of the pancreas, or by reduced sensitivity of tissue cells to insulin. This prevents glucose from being absorbed by cells, causing high levels of blood glucose, or hyperglycemia (high sugar). High blood glucose levels make it difficult for the kidneys to recover all the glucose from nascent urine, resulting in glucose being lost in urine. High glucose levels also result in less water being reabsorbed by the kidneys, causing high amounts of urine to be produced; this may result in dehydration. Over time, high blood glucose levels can cause nerve damage to the eyes and peripheral body tissues, as well as damage to the kidneys and cardiovascular system. Oversecretion of insulin can cause hypoglycemia, low blood glucose levels. This causes insufficient glucose availability to cells, often leading to muscle weakness, and can sometimes cause unconsciousness or death if left untreated.
When blood glucose levels decline below normal levels, for example between meals or when glucose is utilized rapidly during exercise, the hormone glucagon is released from the alpha cells of the pancreas. Glucagon raises blood glucose levels, eliciting what is called a hyperglycemic effect, by stimulating the breakdown of glycogen to glucose in skeletal muscle cells and liver cells in a process called glycogenolysis. Glucose can then be utilized as energy by muscle cells and released into circulation by the liver cells. Glucagon also stimulates absorption of amino acids from the blood by the liver, which then converts them to glucose. This process of glucose synthesis is called gluconeogenesis. Glucagon also stimulates adipose cells to release fatty acids into the blood. These actions mediated by glucagon result in an increase in blood glucose levels to normal homeostatic levels. Rising blood glucose levels inhibit further glucagon release by the pancreas via a negative feedback mechanism. In this way, insulin and glucagon work together to maintain homeostatic glucose levels, as shown in Figure \(5\).
Exercise
Pancreatic tumors may cause excess secretion of glucagon. Type I diabetes results from the failure of the pancreas to produce insulin. Which of the following statement about these two conditions is true?
1. A pancreatic tumor and type I diabetes will have the opposite effects on blood sugar levels.
2. A pancreatic tumor and type I diabetes will both cause hyperglycemia.
3. A pancreatic tumor and type I diabetes will both cause hypoglycemia.
4. Both pancreatic tumors and type I diabetes result in the inability of cells to take up glucose.
Answer
B
Regulation of Blood Glucose Levels by Thyroid Hormones
The basal metabolic rate, which is the amount of calories required by the body at rest, is determined by two hormones produced by the thyroid gland: thyroxine, also known as tetraiodothyronine or T4, and triiodothyronine, also known as T3. These hormones affect nearly every cell in the body except for the adult brain, uterus, testes, blood cells, and spleen. They are transported across the plasma membrane of target cells and bind to receptors on the mitochondria resulting in increased ATP production. In the nucleus, T3 and T4 activate genes involved in energy production and glucose oxidation. This results in increased rates of metabolism and body heat production, which is known as the hormone’s calorigenic effect.
T3 and T4 release from the thyroid gland is stimulated by thyroid-stimulating hormone (TSH), which is produced by the anterior pituitary. TSH binding at the receptors of the follicle of the thyroid triggers the production of T3 and T4 from a glycoprotein called thyroglobulin. Thyroglobulin is present in the follicles of the thyroid, and is converted into thyroid hormones with the addition of iodine. Iodine is formed from iodide ions that are actively transported into the thyroid follicle from the bloodstream. A peroxidase enzyme then attaches the iodine to the tyrosine amino acid found in thyroglobulin. T3 has three iodine ions attached, while T4 has four iodine ions attached. T3 and T4 are then released into the bloodstream, with T4 being released in much greater amounts than T3. As T3 is more active than T4 and is responsible for most of the effects of thyroid hormones, tissues of the body convert T4 to T3 by the removal of an iodine ion. Most of the released T3 and T4 becomes attached to transport proteins in the bloodstream and is unable to cross the plasma membrane of cells. These protein-bound molecules are only released when blood levels of the unattached hormone begin to decline. In this way, a week’s worth of reserve hormone is maintained in the blood. Increased T3 and T4 levels in the blood inhibit the release of TSH, which results in lower T3 and T4 release from the thyroid.
The follicular cells of the thyroid require iodides (anions of iodine) in order to synthesize T3 and T4. Iodides obtained from the diet are actively transported into follicle cells resulting in a concentration that is approximately 30 times higher than in blood. The typical diet in North America provides more iodine than required due to the addition of iodide to table salt. Inadequate iodine intake, which occurs in many developing countries, results in an inability to synthesize T3 and T4 hormones. The thyroid gland enlarges in a condition called goiter, which is caused by overproduction of TSH without the formation of thyroid hormone. Thyroglobulin is contained in a fluid called colloid, and TSH stimulation results in higher levels of colloid accumulation in the thyroid. In the absence of iodine, this is not converted to thyroid hormone, and colloid begins to accumulate more and more in the thyroid gland, leading to goiter.
Disorders can arise from both the underproduction and overproduction of thyroid hormones. Hypothyroidism, underproduction of the thyroid hormones, can cause a low metabolic rate leading to weight gain, sensitivity to cold, and reduced mental activity, among other symptoms. In children, hypothyroidism can cause cretinism, which can lead to mental retardation and growth defects. Hyperthyroidism, the overproduction of thyroid hormones, can lead to an increased metabolic rate and its effects: weight loss, excess heat production, sweating, and an increased heart rate. Graves’ disease is one example of a hyperthyroid condition.
Hormonal Control of Blood Calcium Levels
Regulation of blood calcium concentrations is important for generation of muscle contractions and nerve impulses, which are electrically stimulated. If calcium levels get too high, membrane permeability to sodium decreases and membranes become less responsive. If calcium levels get too low, membrane permeability to sodium increases and convulsions or muscle spasms can result.
Blood calcium levels are regulated by parathyroid hormone (PTH), which is produced by the parathyroid glands, as illustrated in Figure \(6\). PTH is released in response to low blood Ca2+ levels. PTH increases Ca2+ levels by targeting the skeleton, the kidneys, and the intestine. In the skeleton, PTH stimulates osteoclasts, which causes bone to be reabsorbed, releasing Ca2+ from bone into the blood. PTH also inhibits osteoblasts, reducing Ca2+ deposition in bone. In the intestines, PTH increases dietary Ca2+ absorption, and in the kidneys, PTH stimulates reabsorption of the CA2+. While PTH acts directly on the kidneys to increase Ca2+ reabsorption, its effects on the intestine are indirect. PTH triggers the formation of calcitriol, an active form of vitamin D, which acts on the intestines to increase absorption of dietary calcium. PTH release is inhibited by rising blood calcium levels.
Hyperparathyroidism results from an overproduction of parathyroid hormone. This results in excessive calcium being removed from bones and introduced into blood circulation, producing structural weakness of the bones, which can lead to deformation and fractures, plus nervous system impairment due to high blood calcium levels. Hypoparathyroidism, the underproduction of PTH, results in extremely low levels of blood calcium, which causes impaired muscle function and may result in tetany (severe sustained muscle contraction).
The hormone calcitonin, which is produced by the parafollicular or C cells of the thyroid, has the opposite effect on blood calcium levels as does PTH. Calcitonin decreases blood calcium levels by inhibiting osteoclasts, stimulating osteoblasts, and stimulating calcium excretion by the kidneys. This results in calcium being added to the bones to promote structural integrity. Calcitonin is most important in children (when it stimulates bone growth), during pregnancy (when it reduces maternal bone loss), and during prolonged starvation (because it reduces bone mass loss). In healthy nonpregnant, unstarved adults, the role of calcitonin is unclear.
Hormonal Regulation of Growth
Hormonal regulation is required for the growth and replication of most cells in the body. Growth hormone (GH), produced by the anterior portion of the pituitary gland, accelerates the rate of protein synthesis, particularly in skeletal muscle and bones. Growth hormone has direct and indirect mechanisms of action. The first direct action of GH is stimulation of triglyceride breakdown (lipolysis) and release into the blood by adipocytes. This results in a switch by most tissues from utilizing glucose as an energy source to utilizing fatty acids. This process is called a glucose-sparing effect. In another direct mechanism, GH stimulates glycogen breakdown in the liver; the glycogen is then released into the blood as glucose. Blood glucose levels increase as most tissues are utilizing fatty acids instead of glucose for their energy needs. The GH mediated increase in blood glucose levels is called a diabetogenic effect because it is similar to the high blood glucose levels seen in diabetes mellitus.
The indirect mechanism of GH action is mediated by insulin-like growth factors (IGFs) or somatomedins, which are a family of growth-promoting proteins produced by the liver, which stimulates tissue growth. IGFs stimulate the uptake of amino acids from the blood, allowing the formation of new proteins, particularly in skeletal muscle cells, cartilage cells, and other target cells, as shown in Figure \(7\). This is especially important after a meal, when glucose and amino acid concentration levels are high in the blood. GH levels are regulated by two hormones produced by the hypothalamus. GH release is stimulated by growth hormone-releasing hormone (GHRH) and is inhibited by growth hormone-inhibiting hormone (GHIH), also called somatostatin.
A balanced production of growth hormone is critical for proper development. Underproduction of GH in adults does not appear to cause any abnormalities, but in children it can result in pituitary dwarfism, in which growth is reduced. Pituitary dwarfism is characterized by symmetric body formation. In some cases, individuals are under 30 inches in height. Oversecretion of growth hormone can lead to gigantism in children, causing excessive growth. In some documented cases, individuals can reach heights of over eight feet. In adults, excessive GH can lead to acromegaly, a condition in which there is enlargement of bones in the face, hands, and feet that are still capable of growth.
Hormonal Regulation of Stress
When a threat or danger is perceived, the body responds by releasing hormones that will ready it for the “fight-or-flight” response. The effects of this response are familiar to anyone who has been in a stressful situation: increased heart rate, dry mouth, and hair standing up.
Evolution Connection: Fight-or-Flight Response
Interactions of the endocrine hormones have evolved to ensure the body’s internal environment remains stable. Stressors are stimuli that disrupt homeostasis. The sympathetic division of the vertebrate autonomic nervous system has evolved the fight-or-flight response to counter stress-induced disruptions of homeostasis. In the initial alarm phase, the sympathetic nervous system stimulates an increase in energy levels through increased blood glucose levels. This prepares the body for physical activity that may be required to respond to stress: to either fight for survival or to flee from danger.
However, some stresses, such as illness or injury, can last for a long time. Glycogen reserves, which provide energy in the short-term response to stress, are exhausted after several hours and cannot meet long-term energy needs. If glycogen reserves were the only energy source available, neural functioning could not be maintained once the reserves became depleted due to the nervous system’s high requirement for glucose. In this situation, the body has evolved a response to counter long-term stress through the actions of the glucocorticoids, which ensure that long-term energy requirements can be met. The glucocorticoids mobilize lipid and protein reserves, stimulate gluconeogenesis, conserve glucose for use by neural tissue, and stimulate the conservation of salts and water. The mechanisms to maintain homeostasis that are described here are those observed in the human body. However, the fight-or-flight response exists in some form in all vertebrates.
The sympathetic nervous system regulates the stress response via the hypothalamus. Stressful stimuli cause the hypothalamus to signal the adrenal medulla (which mediates short-term stress responses) via nerve impulses, and the adrenal cortex, which mediates long-term stress responses, via the hormone adrenocorticotropic hormone (ACTH), which is produced by the anterior pituitary.
Short-term Stress Response
When presented with a stressful situation, the body responds by calling for the release of hormones that provide a burst of energy. The hormones epinephrine (also known as adrenaline) and norepinephrine (also known as noradrenaline) are released by the adrenal medulla. How do these hormones provide a burst of energy? Epinephrine and norepinephrine increase blood glucose levels by stimulating the liver and skeletal muscles to break down glycogen and by stimulating glucose release by liver cells. Additionally, these hormones increase oxygen availability to cells by increasing the heart rate and dilating the bronchioles. The hormones also prioritize body function by increasing blood supply to essential organs such as the heart, brain, and skeletal muscles, while restricting blood flow to organs not in immediate need, such as the skin, digestive system, and kidneys. Epinephrine and norepinephrine are collectively called catecholamines.
Link to Learning
Watch this Discovery Channel animation describing the flight-or-flight response.
Long-term Stress Response
Long-term stress response differs from short-term stress response. The body cannot sustain the bursts of energy mediated by epinephrine and norepinephrine for long times. Instead, other hormones come into play. In a long-term stress response, the hypothalamus triggers the release of ACTH from the anterior pituitary gland. The adrenal cortex is stimulated by ACTH to release steroid hormones called corticosteroids. Corticosteroids turn on transcription of certain genes in the nuclei of target cells. They change enzyme concentrations in the cytoplasm and affect cellular metabolism. There are two main corticosteroids: glucocorticoids such as cortisol, and mineralocorticoids such as aldosterone. These hormones target the breakdown of fat into fatty acids in the adipose tissue. The fatty acids are released into the bloodstream for other tissues to use for ATP production. The glucocorticoids primarily affect glucose metabolism by stimulating glucose synthesis. Glucocorticoids also have anti-inflammatory properties through inhibition of the immune system. For example, cortisone is used as an anti-inflammatory medication; however, it cannot be used long term as it increases susceptibility to disease due to its immune-suppressing effects.
Mineralocorticoids function to regulate ion and water balance of the body. The hormone aldosterone stimulates the reabsorption of water and sodium ions in the kidney, which results in increased blood pressure and volume.
Hypersecretion of glucocorticoids can cause a condition known as Cushing’s disease, characterized by a shifting of fat storage areas of the body. This can cause the accumulation of adipose tissue in the face and neck, and excessive glucose in the blood. Hyposecretion of the corticosteroids can cause Addison’s disease, which may result in bronzing of the skin, hypoglycemia, and low electrolyte levels in the blood.
Summary
Water levels in the body are controlled by antidiuretic hormone (ADH), which is produced in the hypothalamus and triggers the reabsorption of water by the kidneys. Underproduction of ADH can cause diabetes insipidus. Aldosterone, a hormone produced by the adrenal cortex of the kidneys, enhances Na+ reabsorption from the extracellular fluids and subsequent water reabsorption by diffusion. The renin-angiotensin-aldosterone system is one way that aldosterone release is controlled.
The reproductive system is controlled by the gonadotropins follicle-stimulating hormone (FSH) and luteinizing hormone (LH), which are produced by the pituitary gland. Gonadotropin release is controlled by the hypothalamic hormone gonadotropin-releasing hormone (GnRH). FSH stimulates the maturation of sperm cells in males and is inhibited by the hormone inhibin, while LH stimulates the production of the androgen testosterone. FSH stimulates egg maturation in females, while LH stimulates the production of estrogens and progesterone. Estrogens are a group of steroid hormones produced by the ovaries that trigger the development of secondary sex characteristics in females as well as control the maturation of the ova. In females, the pituitary also produces prolactin, which stimulates milk production after childbirth, and oxytocin, which stimulates uterine contraction during childbirth and milk let-down during suckling.
Insulin is produced by the pancreas in response to rising blood glucose levels and allows cells to utilize blood glucose and store excess glucose for later use. Diabetes mellitus is caused by reduced insulin activity and causes high blood glucose levels, or hyperglycemia. Glucagon is released by the pancreas in response to low blood glucose levels and stimulates the breakdown of glycogen into glucose, which can be used by the body. The body’s basal metabolic rate is controlled by the thyroid hormones thyroxine (T4) and triiodothyronine (T3). The anterior pituitary produces thyroid stimulating hormone (TSH), which controls the release of T3 and T4 from the thyroid gland. Iodine is necessary in the production of thyroid hormone, and the lack of iodine can lead to a condition called goiter.
Parathyroid hormone (PTH) is produced by the parathyroid glands in response to low blood Ca2+ levels. The parafollicular cells of the thyroid produce calcitonin, which reduces blood Ca2+ levels. Growth hormone (GH) is produced by the anterior pituitary and controls the growth rate of muscle and bone. GH action is indirectly mediated by insulin-like growth factors (IGFs). Short-term stress causes the hypothalamus to trigger the adrenal medulla to release epinephrine and norepinephrine, which trigger the fight or flight response. Long-term stress causes the hypothalamus to trigger the anterior pituitary to release adrenocorticotropic hormone (ACTH), which causes the release of corticosteroids, glucocorticoids, and mineralocorticoids, from the adrenal cortex.
Glossary
acromegaly
condition caused by overproduction of GH in adults
Addison’s disease
disorder caused by the hyposecretion of corticosteroids
adrenocorticotropic hormone (ACTH)
hormone released by the anterior pituitary, which stimulates the adrenal cortex to release corticosteroids during the long-term stress response
aldosterone
steroid hormone produced by the adrenal cortex that stimulates the reabsorption of Na+ from extracellular fluids and secretion of K+.
androgen
male sex hormone such as testosterone
antidiuretic hormone (ADH)
hormone produced by the hypothalamus and released by the posterior pituitary that increases water reabsorption by the kidneys
calcitonin
hormone produced by the parafollicular cells of the thyroid gland that functions to lower blood Ca2+ levels and promote bone growth
corticosteroid
hormone released by the adrenal cortex in response to long-term stress
cortisol
glucocorticoid produced in response to stress
Cushing’s disease
disorder caused by the hypersecretion of glucocorticoids
diabetes insipidus
disorder caused by underproduction of ADH
diabetes mellitus
disorder caused by low levels of insulin activity
diabetogenic effect
effect of GH that causes blood glucose levels to rise similar to diabetes mellitus
epinephrine
hormone released by the adrenal medulla in response to a short term stress
estrogens
- a group of steroid hormones, including estradiol and several others, that are produced by the ovaries and elicit secondary sex characteristics in females as well as control the maturation of the ova
follicle-stimulating hormone (FSH)
hormone produced by the anterior pituitary that stimulates gamete production
gigantism
condition caused by overproduction of GH in children
glucagon
hormone produced by the alpha cells of the pancreas in response to low blood sugar; functions to raise blood sugar levels
glucocorticoid
corticosteroid that affects glucose metabolism
gluconeogenesis
synthesis of glucose from amino acids
glucose-sparing effect
effect of GH that causes tissues to use fatty acids instead of glucose as an energy source
glycogenolysis
breakdown of glycogen into glucose
goiter
enlargement of the thyroid gland caused by insufficient dietary iodine levels
gonadotropin
hormone that regulates the gonads, including FSH and LH
growth hormone (GH)
hormone produced by the anterior pituitary that promotes protein synthesis and body growth
growth hormone-inhibiting hormone (GHIH)
hormone produced by the hypothalamus that inhibits growth hormone production, also called somatostatin
growth hormone-releasing hormone (GHRH)
hormone released by the hypothalamus that triggers the release of GH
hyperglycemia
high blood sugar level
hyperthyroidism
overactivity of the thyroid gland
hypoglycemia
low blood sugar level
hypothyroidism
underactivity of the thyroid gland
insulin
hormone produced by the beta cells of the pancreas in response to high blood glucose levels; functions to lower blood glucose levels
insulin-like growth factor (IGF)
growth-promoting protein produced by the liver
mineralocorticoid
corticosteroid that affects ion and water balance
norepinephrine
hormone released by the adrenal medulla in response to a short-term stress hormone production by the gonads
osmoreceptor
receptor in the hypothalamus that monitors the concentration of electrolytes in the blood
oxytocin
hormone released by the posterior pituitary to stimulate uterine contractions during childbirth and milk let-down in the mammary glands
parathyroid hormone (PTH)
hormone produced by the parathyroid glands in response to low blood Ca2+ levels; functions to raise blood Ca2+ levels
pituitary dwarfism
condition caused by underproduction of GH in children
prolactin (PRL)
hormone produced by the anterior pituitary that stimulates milk production
prolactin-inhibiting hormone
hormone produced by the hypothalamus that inhibits the release of prolactin
prolactin-releasing hormone
hormone produced by the hypothalamus that stimulates the release of prolactin
renin
enzyme produced by the juxtaglomerular apparatus of the kidneys that reacts with angiotensinogen to cause the release of aldosterone
thyroglobulin
glycoprotein found in the thyroid that is converted into thyroid hormone
thyroid-stimulating hormone (TSH)
hormone produced by the anterior pituitary that controls the release of T3 and T4 from the thyroid gland
thyroxine (tetraiodothyronine, T4)
thyroid hormone that controls the basal metabolic rate
triiodothyronine (T3)
thyroid hormone that controls the basal metabolic rate | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.04%3A_The_Major_Peripheral_Endocrine_Glands.txt |
Skills to Develop
• Describe the role of different glands in the endocrine system
• Explain how the different glands work together to maintain homeostasis
Both the endocrine and nervous systems use chemical signals to communicate and regulate the body's physiology. The endocrine system releases hormones that act on target cells to regulate development, growth, energy metabolism, reproduction, and many behaviors. The nervous system releases neurotransmitters or neurohormones that regulate neurons, muscle cells, and endocrine cells. Because the neurons can regulate the release of hormones, the nervous and endocrine systems work in a coordinated manner to regulate the body's physiology.
Hypothalamic-Pituitary Axis
The hypothalamus in vertebrates integrates the endocrine and nervous systems. The hypothalamus is an endocrine organ located in the diencephalon of the brain. It receives input from the body and other brain areas and initiates endocrine responses to environmental changes. The hypothalamus acts as an endocrine organ, synthesizing hormones and transporting them along axons to the posterior pituitary gland. It synthesizes and secretes regulatory hormones that control the endocrine cells in the anterior pituitary gland. The hypothalamus contains autonomic centers that control endocrine cells in the adrenal medulla via neuronal control.
The pituitary gland, sometimes called the hypophysis or “master gland” is located at the base of the brain in the sella turcica, a groove of the sphenoid bone of the skull, illustrated in Figure \(1\). It is attached to the hypothalamus via a stalk called the pituitary stalk (or infundibulum). The anterior portion of the pituitary gland is regulated by releasing or release-inhibiting hormones produced by the hypothalamus, and the posterior pituitary receives signals via neurosecretory cells to release hormones produced by the hypothalamus. The pituitary has two distinct regions—the anterior pituitary and the posterior pituitary—which between them secrete nine different peptide or protein hormones. The posterior lobe of the pituitary gland contains axons of the hypothalamic neurons.
Anterior Pituitary
The anterior pituitary gland, or adenohypophysis, is surrounded by a capillary network that extends from the hypothalamus, down along the infundibulum, and to the anterior pituitary. This capillary network is a part of the hypophyseal portal system that carries substances from the hypothalamus to the anterior pituitary and hormones from the anterior pituitary into the circulatory system. A portal system carries blood from one capillary network to another; therefore, the hypophyseal portal system allows hormones produced by the hypothalamus to be carried directly to the anterior pituitary without first entering the circulatory system.
The anterior pituitary produces seven hormones: growth hormone (GH), prolactin (PRL), thyroid-stimulating hormone (TSH), melanin-stimulating hormone (MSH), adrenocorticotropic hormone (ACTH), follicle-stimulating hormone (FSH), and luteinizing hormone (LH). Anterior pituitary hormones are sometimes referred to as tropic hormones, because they control the functioning of other organs. While these hormones are produced by the anterior pituitary, their production is controlled by regulatory hormones produced by the hypothalamus. These regulatory hormones can be releasing hormones or inhibiting hormones, causing more or less of the anterior pituitary hormones to be secreted. These travel from the hypothalamus through the hypophyseal portal system to the anterior pituitary where they exert their effect. Negative feedback then regulates how much of these regulatory hormones are released and how much anterior pituitary hormone is secreted.
Posterior Pituitary
The posterior pituitary is significantly different in structure from the anterior pituitary. It is a part of the brain, extending down from the hypothalamus, and contains mostly nerve fibers and neuroglial cells, which support axons that extend from the hypothalamus to the posterior pituitary. The posterior pituitary and the infundibulum together are referred to as the neurohypophysis.
The hormones antidiuretic hormone (ADH), also known as vasopressin, and oxytocin are produced by neurons in the hypothalamus and transported within these axons along the infundibulum to the posterior pituitary. They are released into the circulatory system via neural signaling from the hypothalamus. These hormones are considered to be posterior pituitary hormones, even though they are produced by the hypothalamus, because that is where they are released into the circulatory system. The posterior pituitary itself does not produce hormones, but instead stores hormones produced by the hypothalamus and releases them into the blood stream.
Thyroid Gland
The thyroid gland is located in the neck, just below the larynx and in front of the trachea, as shown in Figure \(2\). It is a butterfly-shaped gland with two lobes that are connected by the isthmus. It has a dark red color due to its extensive vascular system. When the thyroid swells due to dysfunction, it can be felt under the skin of the neck.
The thyroid gland is made up of many spherical thyroid follicles, which are lined with a simple cuboidal epithelium. These follicles contain a viscous fluid, called colloid, which stores the glycoprotein thyroglobulin, the precursor to the thyroid hormones. The follicles produce hormones that can be stored in the colloid or released into the surrounding capillary network for transport to the rest of the body via the circulatory system.
Thyroid follicle cells synthesize the hormone thyroxine, which is also known as T4 because it contains four atoms of iodine, and triiodothyronine, also known as T3 because it contains three atoms of iodine. Follicle cells are stimulated to release stored T3 and T4 by thyroid stimulating hormone (TSH), which is produced by the anterior pituitary. These thyroid hormones increase the rates of mitochondrial ATP production.
A third hormone, calcitonin, is produced by parafollicular cells of the thyroid either releasing hormones or inhibiting hormones. Calcitonin release is not controlled by TSH, but instead is released when calcium ion concentrations in the blood rise. Calcitonin functions to help regulate calcium concentrations in body fluids. It acts in the bones to inhibit osteoclast activity and in the kidneys to stimulate excretion of calcium. The combination of these two events lowers body fluid levels of calcium.
Parathyroid Glands
Most people have four parathyroid glands; however, the number can vary from two to six. These glands are located on the posterior surface of the thyroid gland, as shown in Figure \(3\). Normally, there is a superior gland and an inferior gland associated with each of the thyroid’s two lobes. Each parathyroid gland is covered by connective tissue and contains many secretory cells that are associated with a capillary network.
The parathyroid glands produce parathyroid hormone (PTH). PTH increases blood calcium concentrations when calcium ion levels fall below normal. PTH (1) enhances reabsorption of Ca2+ by the kidneys, (2) stimulates osteoclast activity and inhibits osteoblast activity, and (3) it stimulates synthesis and secretion of calcitriol by the kidneys, which enhances Ca2+ absorption by the digestive system. PTH is produced by chief cells of the parathyroid. PTH and calcitonin work in opposition to one another to maintain homeostatic Ca2+ levels in body fluids. Another type of cells, oxyphil cells, exist in the parathyroid but their function is not known. These hormones encourage bone growth, muscle mass, and blood cell formation in children and women.
Adrenal Glands
The adrenal glands are associated with the kidneys; one gland is located on top of each kidney as illustrated in Figure \(4\). The adrenal glands consist of an outer adrenal cortex and an inner adrenal medulla. These regions secrete different hormones.
Adrenal Cortex
The adrenal cortex is made up of layers of epithelial cells and associated capillary networks. These layers form three distinct regions: an outer zona glomerulosa that produces mineralocorticoids, a middle zona fasciculata that produces glucocorticoids, and an inner zona reticularis that produces androgens.
The main mineralocorticoid is aldosterone, which regulates the concentration of Na+ ions in urine, sweat, pancreas, and saliva. Aldosterone release from the adrenal cortex is stimulated by a decrease in blood concentrations of sodium ions, blood volume, or blood pressure, or by an increase in blood potassium levels.
The three main glucocorticoids are cortisol, corticosterone, and cortisone. The glucocorticoids stimulate the synthesis of glucose and gluconeogenesis (converting a non-carbohydrate to glucose) by liver cells and they promote the release of fatty acids from adipose tissue. These hormones increase blood glucose levels to maintain levels within a normal range between meals. These hormones are secreted in response to ACTH and levels are regulated by negative feedback.
Androgens are sex hormones that promote masculinity. They are produced in small amounts by the adrenal cortex in both males and females. They do not affect sexual characteristics and may supplement sex hormones released from the gonads.
Adrenal Medulla
The adrenal medulla contains large, irregularly shaped cells that are closely associated with blood vessels. These cells are innervated by preganglionic autonomic nerve fibers from the central nervous system.
The adrenal medulla contains two types of secretory cells: one that produces epinephrine (adrenaline) and another that produces norepinephrine (noradrenaline). Epinephrine is the primary adrenal medulla hormone accounting for 75 to 80 percent of its secretions. Epinephrine and norepinephrine increase heart rate, breathing rate, cardiac muscle contractions, blood pressure, and blood glucose levels. They also accelerate the breakdown of glucose in skeletal muscles and stored fats in adipose tissue.
The release of epinephrine and norepinephrine is stimulated by neural impulses from the sympathetic nervous system. Secretion of these hormones is stimulated by acetylcholine release from preganglionic sympathetic fibers innervating the adrenal medulla. These neural impulses originate from the hypothalamus in response to stress to prepare the body for the fight-or-flight response.
Pancreas
The pancreas, illustrated in Figure \(5\), is an elongated organ that is located between the stomach and the proximal portion of the small intestine. It contains both exocrine cells that excrete digestive enzymes and endocrine cells that release hormones. It is sometimes referred to as a heterocrine gland because it has both endocrine and exocrine functions.
The endocrine cells of the pancreas form clusters called pancreatic islets or the islets of Langerhans, as visible in the micrograph shown in Figure \(6\). The pancreatic islets contain two primary cell types: alpha cells, which produce the hormone glucagon, and beta cells, which produce the hormone insulin. These hormones regulate blood glucose levels. As blood glucose levels decline, alpha cells release glucagon to raise the blood glucose levels by increasing rates of glycogen breakdown and glucose release by the liver. When blood glucose levels rise, such as after a meal, beta cells release insulin to lower blood glucose levels by increasing the rate of glucose uptake in most body cells, and by increasing glycogen synthesis in skeletal muscles and the liver. Together, glucagon and insulin regulate blood glucose levels.
Pineal Gland
The pineal gland produces melatonin. The rate of melatonin production is affected by the photoperiod. Collaterals from the visual pathways innervate the pineal gland. During the day photoperiod, little melatonin is produced; however, melatonin production increases during the dark photoperiod (night). In some mammals, melatonin has an inhibitory affect on reproductive functions by decreasing production and maturation of sperm, oocytes, and reproductive organs. Melatonin is an effective antioxidant, protecting the CNS from free radicals such as nitric oxide and hydrogen peroxide. Lastly, melatonin is involved in biological rhythms, particularly circadian rhythms such as the sleep-wake cycle and eating habits.
Gonads
The gonads—the male testes and female ovaries—produce steroid hormones. The testes produce androgens, testosterone being the most prominent, which allow for the development of secondary sex characteristics and the production of sperm cells. The ovaries produce estradiol and progesterone, which cause secondary sex characteristics and prepare the body for childbirth.
Table \(1\): Endocrine Glands and their Associated Hormones
Endocrine Gland Associated Hormones Effect
Hypothalamus releasing and inhibiting hormones regulate hormone release from pituitary gland; produce oxytocin; produce uterine contractions and milk secretion in females
antidiuretic hormone (ADH) water reabsorption from kidneys; vasoconstriction to increase blood pressure
Pituitary (Anterior) growth hormone (GH) promotes growth of body tissues, protein synthesis; metabolic functions
prolactin (PRL) promotes milk production
thyroid stimulating hormone (TSH) stimulates thyroid hormone release
adrenocorticotropic hormone (ACTH) stimulates hormone release by adrenal cortex, glucocorticoids
follicle-stimulating hormone (FSH) stimulates gamete production (both ova and sperm); secretion of estradiol
luteinizing hormone (LH) stimulates androgen production by gonads; ovulation, secretion of progesterone
melanocyte-stimulating hormone (MSH) stimulates melanocytes of the skin increasing melanin pigment production.
Pituitary (Posterior) antidiuretic hormone (ADH) stimulates water reabsorption by kidneys
oxytocin stimulates uterine contractions during childbirth; milk ejection; stimulates ductus deferens and prostate gland contraction during emission
Thyroid thyroxine, triiodothyronine stimulate and maintain metabolism; growth and development
calcitonin reduces blood Ca2+ levels
Parathyroid parathyroid hormone (PTH) increases blood Ca2+ levels
Adrenal (Cortex) aldosterone increases blood Na+ levels; increase K+ secretion
cortisol, corticosterone, cortisone increase blood glucose levels; anti-inflammatory effects
Adrenal (Medulla) epinephrine, norepinephrine stimulate fight-or-flight response; increase blood gluclose levels; increase metabolic activities
Pancreas insulin reduces blood glucose levels
glucagon increases blood glucose levels
Pineal gland melatonin regulates some biological rhythms and protects CNS from free radicals
Testes androgens regulate, promote, increase or maintain sperm production; male secondary sexual characteristics
Ovaries estrogen promotes uterine lining growth; female secondary sexual characteristics
progestins promote and maintain uterine lining growth
Organs with Secondary Endocrine Functions
There are several organs whose primary functions are non-endocrine but that also possess endocrine functions. These include the heart, kidneys, intestines, thymus, gonads, and adipose tissue.
The heart possesses endocrine cells in the walls of the atria that are specialized cardiac muscle cells. These cells release the hormone atrial natriuretic peptide (ANP) in response to increased blood volume. High blood volume causes the cells to be stretched, resulting in hormone release. ANP acts on the kidneys to reduce the reabsorption of Na+, causing Na+ and water to be excreted in the urine. ANP also reduces the amounts of renin released by the kidneys and aldosterone released by the adrenal cortex, further preventing the retention of water. In this way, ANP causes a reduction in blood volume and blood pressure, and reduces the concentration of Na+ in the blood.
The gastrointestinal tract produces several hormones that aid in digestion. The endocrine cells are located in the mucosa of the GI tract throughout the stomach and small intestine. Some of the hormones produced include gastrin, secretin, and cholecystokinin, which are secreted in the presence of food, and some of which act on other organs such as the pancreas, gallbladder, and liver. They trigger the release of gastric juices, which help to break down and digest food in the GI tract.
While the adrenal glands associated with the kidneys are major endocrine glands, the kidneys themselves also possess endocrine function. Renin is released in response to decreased blood volume or pressure and is part of the renin-angiotensin-aldosterone system that leads to the release of aldosterone. Aldosterone then causes the retention of Na+ and water, raising blood volume. The kidneys also release calcitriol, which aids in the absorption of Ca2+ and phosphate ions. Erythropoietin (EPO) is a protein hormone that triggers the formation of red blood cells in the bone marrow. EPO is released in response to low oxygen levels. Because red blood cells are oxygen carriers, increased production results in greater oxygen delivery throughout the body. EPO has been used by athletes to improve performance, as greater oxygen delivery to muscle cells allows for greater endurance. Because red blood cells increase the viscosity of blood, artificially high levels of EPO can cause severe health risks.
The thymus is found behind the sternum; it is most prominent in infants, becoming smaller in size through adulthood. The thymus produces hormones referred to as thymosins, which contribute to the development of the immune response.
Adipose tissue is a connective tissue found throughout the body. It produces the hormone leptin in response to food intake. Leptin increases the activity of anorexigenic neurons and decreases that of orexigenic neurons, producing a feeling of satiety after eating, thus affecting appetite and reducing the urge for further eating. Leptin is also associated with reproduction. It must be present for GnRH and gonadotropin synthesis to occur. Extremely thin females may enter puberty late; however, if adipose levels increase, more leptin will be produced, improving fertility.
Summary
The pituitary gland is located at the base of the brain and is attached to the hypothalamus by the infundibulum. The anterior pituitary receives products from the hypothalamus by the hypophyseal portal system and produces six hormones. The posterior pituitary is an extension of the brain and releases hormones (antidiuretic hormone and oxytocin) produced by the hypothalamus.
The thyroid gland is located in the neck and is composed of two lobes connected by the isthmus. The thyroid is made up of follicle cells that produce the hormones thyroxine and triiodothyronine. Parafollicular cells of the thyroid produce calcitonin. The parathyroid glands lie on the posterior surface of the thyroid gland and produce parathyroid hormone.
The adrenal glands are located on top of the kidneys and consist of the renal cortex and renal medulla. The adrenal cortex is the outer part of the adrenal gland and produces the corticosteroids, glucocorticoids, and mineralocorticoids. The adrenal medulla is the inner part of the adrenal gland and produces the catecholamines epinephrine and norepinephrine.
The pancreas lies in the abdomen between the stomach and the small intestine. Clusters of endocrine cells in the pancreas form the islets of Langerhans, which are composed of alpha cells that release glucagon and beta cells that release insulin.
Some organs possess endocrine activity as a secondary function but have another primary function. The heart produces the hormone atrial natriuretic peptide, which functions to reduce blood volume, pressure, and Na+ concentration. The gastrointestinal tract produces various hormones that aid in digestion. The kidneys produce renin, calcitriol, and erythropoietin. Adipose tissue produces leptin, which promotes satiety signals in the brain.
Glossary
adrenal cortex
outer portion of adrenal glands that produces corticosteroids
adrenal gland
endocrine glands associated with the kidneys
adrenal medulla
inner portion of adrenal glands that produces epinephrine and norepinephrine
alpha cell
endocrine cell of the pancreatic islets that produces the hormone glucagon
anterior pituitary
portion of the pituitary gland that produces six hormones; also called adenohypophysis
atrial natriuretic peptide (ANP)
hormone produced by the heart to reduce blood volume, pressure, and Na+ concentration
beta cell
endocrine cell of the pancreatic islets that produces the hormone insulin
colloid
fluid inside the thyroid gland that contains the glycoprotein thyroglobulin
endocrine gland
gland that secretes hormones into the surrounding interstitial fluid, which then diffuse into blood and are carried to various organs and tissues within the body
erythropoietin (EPO)
hormone produced by the kidneys to stimulate red blood cell production in the bone marrow
hypophyseal portal system
system of blood vessels that carries hormones from the hypothalamus to the anterior pituitary
islets of Langerhans (pancreatic islets)
endocrine cells of the pancreas
isthmus
tissue mass that connects the two lobes of the thyroid gland
leptin
hormone produced by adipose tissue that promotes feelings of satiety and reduces hunger
pancreas
organ located between the stomach and the small intestine that contains exocrine and endocrine cells
parafollicular cell
thyroid cell that produces the hormone calcitonin
parathyroid gland
gland located on the surface of the thyroid that produces parathyroid hormone
pituitary gland
endocrine gland located at the base of the brain composed of an anterior and posterior region; also called hypophysis
pituitary stalk
(also, infundibulum) stalk that connects the pituitary gland to the hypothalamus
posterior pituitary
extension of the brain that releases hormones produced by the hypothalamus; along with the infundibulum, it is also referred to as the neurohypophysis
thymus
gland located behind the sternum that produces thymosin hormones that contribute to the development of the immune system
thyroid gland
endocrine gland located in the neck that produces thyroid hormones thyroxine and triiodothyronine | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/44%3A_The_Endocrine_System/44.05%3A_Other_Hormones_and_Their_Effects.txt |
• 45.1: Types of Skeletal Systems
A skeletal system is necessary to support the body, protect internal organs, and allow for the movement of an organism. There are three different skeleton designs that fulfill these functions: hydrostatic skeleton, exoskeleton, and endoskeleton.
• 45.2: A Closer Look at Bone
Bone, or osseous tissue, is a connective tissue that constitutes the endoskeleton. It contains specialized cells and a matrix of mineral salts and collagen fibers. The bones of the human skeleton are classified by their shape: long bones, short bones, flat bones, sutural bones, sesamoid bones, and irregular bones.
• 45.3: Joints
The point at which two or more bones meet is called a joint, or articulation. Joints are responsible for movement, such as the movement of limbs, and stability, such as the stability found in the bones of the skull.
• 45.4: Muscle Contraction
The body contains three types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle. Skeleton muscle tissue is composed of sarcomeres, the functional units of muscle tissue. Muscle contraction occurs when sarcomeres shorten, as thick and thin filaments slide past each other, which is called the sliding filament model of muscle contraction. ATP provides the energy for cross-bridge formation and filament sliding.
• 45.5: Vertebrate Skeleton Evolution and Modes of Locomotion
A skeletal system is necessary to support the body, protect internal organs, and allow for the movement of an organism. There are three different skeleton designs that fulfill these functions: hydrostatic skeleton, exoskeleton, and endoskeleton.
45: The Musculoskeletal System
Skills to Develop
• Discuss the different types of skeletal systems
• Explain the role of the human skeletal system
• Compare and contrast different skeletal systems
A skeletal system is necessary to support the body, protect internal organs, and allow for the movement of an organism. There are three different skeleton designs that fulfill these functions: hydrostatic skeleton, exoskeleton, and endoskeleton.
Hydrostatic Skeleton
A hydrostatic skeleton is a skeleton formed by a fluid-filled compartment within the body, called the coelom. The organs of the coelom are supported by the aqueous fluid, which also resists external compression. This compartment is under hydrostatic pressure because of the fluid and supports the other organs of the organism. This type of skeletal system is found in soft-bodied animals such as sea anemones, earthworms, Cnidaria, and other invertebrates (Figure \(1\)).
Movement in a hydrostatic skeleton is provided by muscles that surround the coelom. The muscles in a hydrostatic skeleton contract to change the shape of the coelom; the pressure of the fluid in the coelom produces movement. For example, earthworms move by waves of muscular contractions of the skeletal muscle of the body wall hydrostatic skeleton, called peristalsis, which alternately shorten and lengthen the body. Lengthening the body extends the anterior end of the organism. Most organisms have a mechanism to fix themselves in the substrate. Shortening the muscles then draws the posterior portion of the body forward. Although a hydrostatic skeleton is well-suited to invertebrate organisms such as earthworms and some aquatic organisms, it is not an efficient skeleton for terrestrial animals.
Exoskeleton
An exoskeleton is an external skeleton that consists of a hard encasement on the surface of an organism. For example, the shells of crabs and insects are exoskeletons (Figure \(2\)). This skeleton type provides defence against predators, supports the body, and allows for movement through the contraction of attached muscles. As with vertebrates, muscles must cross a joint inside the exoskeleton. Shortening of the muscle changes the relationship of the two segments of the exoskeleton. Arthropods such as crabs and lobsters have exoskeletons that consist of 30–50 percent chitin, a polysaccharide derivative of glucose that is a strong but flexible material. Chitin is secreted by the epidermal cells. The exoskeleton is further strengthened by the addition of calcium carbonate in organisms such as the lobster. Because the exoskeleton is acellular, arthropods must periodically shed their exoskeletons because the exoskeleton does not grow as the organism grows.
Endoskeleton
An endoskeleton is a skeleton that consists of hard, mineralized structures located within the soft tissue of organisms. An example of a primitive endoskeletal structure is the spicules of sponges. The bones of vertebrates are composed of tissues, whereas sponges have no true tissues (Figure \(1\)). Endoskeletons provide support for the body, protect internal organs, and allow for movement through contraction of muscles attached to the skeleton.
The human skeleton is an endoskeleton that consists of 206 bones in the adult. It has five main functions: providing support to the body, storing minerals and lipids, producing blood cells, protecting internal organs, and allowing for movement. The skeletal system in vertebrates is divided into the axial skeleton (which consists of the skull, vertebral column, and rib cage), and the appendicular skeleton (which consists of the shoulders, limb bones, the pectoral girdle, and the pelvic girdle).
Human Axial Skeleton
The axial skeleton forms the central axis of the body and includes the bones of the skull, ossicles of the middle ear, hyoid bone of the throat, vertebral column, and the thoracic cage (ribcage) (Figure \(1\)). The function of the axial skeleton is to provide support and protection for the brain, the spinal cord, and the organs in the ventral body cavity. It provides a surface for the attachment of muscles that move the head, neck, and trunk, performs respiratory movements, and stabilizes parts of the appendicular skeleton.
The Skull
The bones of the skull support the structures of the face and protect the brain. The skull consists of 22 bones, which are divided into two categories: cranial bones and facial bones. The cranial bones are eight bones that form the cranial cavity, which encloses the brain and serves as an attachment site for the muscles of the head and neck. The eight cranial bones are the frontal bone, two parietal bones, two temporal bones, occipital bone, sphenoid bone, and the ethmoid bone. Although the bones developed separately in the embryo and fetus, in the adult, they are tightly fused with connective tissue and adjoining bones do not move (Figure \(5\)).
The auditory ossicles of the middle ear transmit sounds from the air as vibrations to the fluid-filled cochlea. The auditory ossicles consist of six bones: two malleus bones, two incus bones, and two stapes on each side. These are the smallest bones in the body and are unique to mammals.
Fourteen facial bones form the face, provide cavities for the sense organs (eyes, mouth, and nose), protect the entrances to the digestive and respiratory tracts, and serve as attachment points for facial muscles. The 14 facial bones are the nasal bones, the maxillary bones, zygomatic bones, palatine, vomer, lacrimal bones, the inferior nasal conchae, and the mandible. All of these bones occur in pairs except for the mandible and the vomer (Figure \(6\)).
Although it is not found in the skull, the hyoid bone is considered a component of the axial skeleton. The hyoid bone lies below the mandible in the front of the neck. It acts as a movable base for the tongue and is connected to muscles of the jaw, larynx, and tongue. The mandible articulates with the base of the skull. The mandible controls the opening to the airway and gut. In animals with teeth, the mandible brings the surfaces of the teeth in contact with the maxillary teeth.
The Vertebral Column
The vertebral column, or spinal column, surrounds and protects the spinal cord, supports the head, and acts as an attachment point for the ribs and muscles of the back and neck. The adult vertebral column comprises 26 bones: the 24 vertebrae, the sacrum, and the coccyx bones. In the adult, the sacrum is typically composed of five vertebrae that fuse into one. The coccyx is typically 3–4 vertebrae that fuse into one. Around the age of 70, the sacrum and the coccyx may fuse together. We begin life with approximately 33 vertebrae, but as we grow, several vertebrae fuse together. The adult vertebrae are further divided into the 7 cervical vertebrae, 12 thoracic vertebrae, and 5 lumbar vertebrae (Figure \(7\)).
Each vertebral body has a large hole in the center through which the nerves of the spinal cord pass. There is also a notch on each side through which the spinal nerves, which serve the body at that level, can exit from the spinal cord. The vertebral column is approximately 71 cm (28 inches) in adult male humans and is curved, which can be seen from a side view. The names of the spinal curves correspond to the region of the spine in which they occur. The thoracic and sacral curves are concave (curve inwards relative to the front of the body) and the cervical and lumbar curves are convex (curve outwards relative to the front of the body). The arched curvature of the vertebral column increases its strength and flexibility, allowing it to absorb shocks like a spring (Figure \(7\)).
Intervertebral discs composed of fibrous cartilage lie between adjacent vertebral bodies from the second cervical vertebra to the sacrum. Each disc is part of a joint that allows for some movement of the spine and acts as a cushion to absorb shocks from movements such as walking and running. Intervertebral discs also act as ligaments to bind vertebrae together. The inner part of discs, the nucleus pulposus, hardens as people age and becomes less elastic. This loss of elasticity diminishes its ability to absorb shocks.
The Thoracic Cage
The thoracic cage, also known as the ribcage, is the skeleton of the chest, and consists of the ribs, sternum, thoracic vertebrae, and costal cartilages (Figure \(8\).8). The thoracic cage encloses and protects the organs of the thoracic cavity, including the heart and lungs. It also provides support for the shoulder girdles and upper limbs, and serves as the attachment point for the diaphragm, muscles of the back, chest, neck, and shoulders. Changes in the volume of the thorax enable breathing.
The sternum, or breastbone, is a long, flat bone located at the anterior of the chest. It is formed from three bones that fuse in the adult. The ribs are 12 pairs of long, curved bones that attach to the thoracic vertebrae and curve toward the front of the body, forming the ribcage. Costal cartilages connect the anterior ends of the ribs to the sternum, with the exception of rib pairs 11 and 12, which are free-floating ribs.
Human Appendicular Skeleton
The appendicular skeleton is composed of the bones of the upper limbs (which function to grasp and manipulate objects) and the lower limbs (which permit locomotion). It also includes the pectoral girdle, or shoulder girdle, that attaches the upper limbs to the body, and the pelvic girdle that attaches the lower limbs to the body (Figure \(9\)).
The Pectoral Girdle
The pectoral girdle bones provide the points of attachment of the upper limbs to the axial skeleton. The human pectoral girdle consists of the clavicle (or collarbone) in the anterior, and the scapula (or shoulder blades) in the posterior (Figure \(10\)).
The clavicles are S-shaped bones that position the arms on the body. The clavicles lie horizontally across the front of the thorax (chest) just above the first rib. These bones are fairly fragile and are susceptible to fractures. For example, a fall with the arms outstretched causes the force to be transmitted to the clavicles, which can break if the force is excessive. The clavicle articulates with the sternum and the scapula.
The scapulae are flat, triangular bones that are located at the back of the pectoral girdle. They support the muscles crossing the shoulder joint. A ridge, called the spine, runs across the back of the scapula and can easily be felt through the skin (Figure \(10\)). The spine of the scapula is a good example of a bony protrusion that facilitates a broad area of attachment for muscles to bone.
The Upper Limb
The upper limb contains 30 bones in three regions: the arm (shoulder to elbow), the forearm (ulna and radius), and the wrist and hand (Figure \(11\)).
An articulation is any place at which two bones are joined. The humerus is the largest and longest bone of the upper limb and the only bone of the arm. It articulates with the scapula at the shoulder and with the forearm at the elbow. The forearm extends from the elbow to the wrist and consists of two bones: the ulna and the radius. The radius is located along the lateral (thumb) side of the forearm and articulates with the humerus at the elbow. The ulna is located on the medial aspect (pinky-finger side) of the forearm. It is longer than the radius. The ulna articulates with the humerus at the elbow. The radius and ulna also articulate with the carpal bones and with each other, which in vertebrates enables a variable degree of rotation of the carpus with respect to the long axis of the limb. The hand includes the eight bones of the carpus (wrist), the five bones of the metacarpus (palm), and the 14 bones of the phalanges (digits). Each digit consists of three phalanges, except for the thumb, when present, which has only two.
The Pelvic Girdle
The pelvic girdle attaches to the lower limbs of the axial skeleton. Because it is responsible for bearing the weight of the body and for locomotion, the pelvic girdle is securely attached to the axial skeleton by strong ligaments. It also has deep sockets with robust ligaments to securely attach the femur to the body. The pelvic girdle is further strengthened by two large hip bones. In adults, the hip bones, or coxal bones, are formed by the fusion of three pairs of bones: the ilium, ischium, and pubis. The pelvis joins together in the anterior of the body at a joint called the pubic symphysis and with the bones of the sacrum at the posterior of the body.
The female pelvis is slightly different from the male pelvis. Over generations of evolution, females with a wider pubic angle and larger diameter pelvic canal reproduced more successfully. Therefore, their offspring also had pelvic anatomy that enabled successful childbirth (Figure \(12\)).
The Lower Limb
The lower limb consists of the thigh, the leg, and the foot. The bones of the lower limb are the femur (thigh bone), patella (kneecap), tibia and fibula (bones of the leg), tarsals (bones of the ankle), and metatarsals and phalanges (bones of the foot) (Figure \(13\)). The bones of the lower limbs are thicker and stronger than the bones of the upper limbs because of the need to support the entire weight of the body and the resulting forces from locomotion. In addition to evolutionary fitness, the bones of an individual will respond to forces exerted upon them.
The femur, or thighbone, is the longest, heaviest, and strongest bone in the body. The femur and pelvis form the hip joint at the proximal end. At the distal end, the femur, tibia, and patella form the knee joint. The patella, or kneecap, is a triangular bone that lies anterior to the knee joint. The patella is embedded in the tendon of the femoral extensors (quadriceps). It improves knee extension by reducing friction. The tibia, or shinbone, is a large bone of the leg that is located directly below the knee. The tibia articulates with the femur at its proximal end, with the fibula and the tarsal bones at its distal end. It is the second largest bone in the human body and is responsible for transmitting the weight of the body from the femur to the foot. The fibula, or calf bone, parallels and articulates with the tibia. It does not articulate with the femur and does not bear weight. The fibula acts as a site for muscle attachment and forms the lateral part of the ankle joint.
The tarsals are the seven bones of the ankle. The ankle transmits the weight of the body from the tibia and the fibula to the foot. The metatarsals are the five bones of the foot. The phalanges are the 14 bones of the toes. Each toe consists of three phalanges, except for the big toe that has only two (Figure \(14\)). Variations exist in other species; for example, the horse’s metacarpals and metatarsals are oriented vertically and do not make contact with the substrate.
Evolution Connection: Evolution of Body Design for Locomotion on Land
The transition of vertebrates onto land required a number of changes in body design, as movement on land presents a number of challenges for animals that are adapted to movement in water. The buoyancy of water provides a certain amount of lift, and a common form of movement by fish is lateral undulations of the entire body. This back and forth movement pushes the body against the water, creating forward movement. In most fish, the muscles of paired fins attach to girdles within the body, allowing for some control of locomotion. As certain fish began moving onto land, they retained their lateral undulation form of locomotion (anguilliform). However, instead of pushing against water, their fins or flippers became points of contact with the ground, around which they rotated their bodies.
The effect of gravity and the lack of buoyancy on land meant that body weight was suspended on the limbs, leading to increased strengthening and ossification of the limbs. The effect of gravity also required changes to the axial skeleton. Lateral undulations of land animal vertebral columns cause torsional strain. A firmer, more ossified vertebral column became common in terrestrial tetrapods because it reduces strain while providing the strength needed to support the body’s weight. In later tetrapods, the vertebrae began allowing for vertical motion rather than lateral flexion. Another change in the axial skeleton was the loss of a direct attachment between the pectoral girdle and the head. This reduced the jarring to the head caused by the impact of the limbs on the ground. The vertebrae of the neck also evolved to allow movement of the head independently of the body.
The appendicular skeleton of land animals is also different from aquatic animals. The shoulders attach to the pectoral girdle through muscles and connective tissue, thus reducing the jarring of the skull. Because of a lateral undulating vertebral column, in early tetrapods, the limbs were splayed out to the side and movement occurred by performing “push-ups.” The vertebrae of these animals had to move side-to-side in a similar manner to fish and reptiles. This type of motion requires large muscles to move the limbs toward the midline; it was almost like walking while doing push-ups, and it is not an efficient use of energy. Later tetrapods have their limbs placed under their bodies, so that each stride requires less force to move forward. This resulted in decreased adductor muscle size and an increased range of motion of the scapulae. This also restricts movement primarily to one plane, creating forward motion rather than moving the limbs upward as well as forward. The femur and humerus were also rotated, so that the ends of the limbs and digits were pointed forward, in the direction of motion, rather than out to the side. By placement underneath the body, limbs can swing forward like a pendulum to produce a stride that is more efficient for moving over land.
Summary
The three types of skeleton designs are hydrostatic skeletons, exoskeletons, and endoskeletons. A hydrostatic skeleton is formed by a fluid-filled compartment held under hydrostatic pressure; movement is created by the muscles producing pressure on the fluid. An exoskeleton is a hard external skeleton that protects the outer surface of an organism and enables movement through muscles attached on the inside. An endoskeleton is an internal skeleton composed of hard, mineralized tissue that also enables movement by attachment to muscles. The human skeleton is an endoskeleton that is composed of the axial and appendicular skeleton. The axial skeleton is composed of the bones of the skull, ossicles of the ear, hyoid bone, vertebral column, and ribcage. The skull consists of eight cranial bones and 14 facial bones. Six bones make up the ossicles of the middle ear, while the hyoid bone is located in the neck under the mandible. The vertebral column contains 26 bones, and it surrounds and protects the spinal cord. The thoracic cage consists of the sternum, ribs, thoracic vertebrae, and costal cartilages. The appendicular skeleton is made up of the limbs of the upper and lower limbs. The pectoral girdle is composed of the clavicles and the scapulae. The upper limb contains 30 bones in the arm, the forearm, and the hand. The pelvic girdle attaches the lower limbs to the axial skeleton. The lower limb includes the bones of the thigh, the leg, and the foot.
Glossary
appendicular skeleton
composed of the bones of the upper limbs, which function to grasp and manipulate objects, and the lower limbs, which permit locomotion
articulation
any place where two bones are joined
auditory ossicle
(also, middle ear) transduces sounds from the air into vibrations in the fluid-filled cochlea
axial skeleton
forms the central axis of the body and includes the bones of the skull, the ossicles of the middle ear, the hyoid bone of the throat, the vertebral column, and the thoracic cage (ribcage)
carpus
eight bones that comprise the wrist
clavicle
S-shaped bone that positions the arms laterally
coxal bone
hip bone
cranial bone
one of eight bones that form the cranial cavity that encloses the brain and serves as an attachment site for the muscles of the head and neck
endoskeleton
skeleton of living cells that produce a hard, mineralized tissue located within the soft tissue of organisms
exoskeleton
a secreted cellular product external skeleton that consists of a hard encasement on the surface of an organism
facial bone
one of the 14 bones that form the face; provides cavities for the sense organs (eyes, mouth, and nose) and attachment points for facial muscles
femur
(also, thighbone) longest, heaviest, and strongest bone in the body
fibula
(also, calf bone) parallels and articulates with the tibia
forearm
extends from the elbow to the wrist and consists of two bones: the ulna and the radius
humerus
only bone of the arm
hydrostatic skeleton
skeleton that consists of aqueous fluid held under pressure in a closed body compartment
hyoid bone
lies below the mandible in the front of the neck
intervertebral disc
composed of fibrous cartilage; lies between adjacent vertebrae from the second cervical vertebra to the sacrum
lower limb
consists of the thigh, the leg, and the foot
metacarpus
five bones that comprise the palm
metatarsal
one of the five bones of the foot
patella
(also, kneecap) triangular bone that lies anterior to the knee joint
pectoral girdle
bones that transmit the force generated by the upper limbs to the axial skeleton
phalange
one of the bones of the fingers or toes
pelvic girdle
bones that transmit the force generated by the lower limbs to the axial skeleton
radius
bone located along the lateral (thumb) side of the forearm; articulates with the humerus at the elbow
rib
one of 12 pairs of long, curved bones that attach to the thoracic vertebrae and curve toward the front of the body to form the ribcage
scapula
flat, triangular bone located at the posterior pectoral girdle
skull
bone that supports the structures of the face and protects the brain
sternum
(also, breastbone) long, flat bone located at the front of the chest
tarsal
one of the seven bones of the ankle
thoracic cage
(also, ribcage) skeleton of the chest, which consists of the ribs, thoracic vertebrae, sternum, and costal cartilages
tibia
(also, shinbone) large bone of the leg that is located directly below the knee
ulna
bone located on the medial aspect (pinky-finger side) of the forearm
vertebral column
(also, spine) surrounds and protects the spinal cord, supports the head, and acts as an attachment point for ribs and muscles of the back and neck | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/45%3A_The_Musculoskeletal_System/45.01%3A_Types_of_Skeletal_Systems.txt |
Skills to Develop
• Classify the different types of bones in the skeleton
• Explain the role of the different cell types in bone
• Explain how bone forms during development
Bone, or osseous tissue, is a connective tissue that constitutes the endoskeleton. It contains specialized cells and a matrix of mineral salts and collagen fibers.
The mineral salts primarily include hydroxyapatite, a mineral formed from calcium phosphate. Calcification is the process of deposition of mineral salts on the collagen fiber matrix that crystallizes and hardens the tissue. The process of calcification only occurs in the presence of collagen fibers.
The bones of the human skeleton are classified by their shape: long bones, short bones, flat bones, sutural bones, sesamoid bones, and irregular bones (Figure \(1\)).
Long bones are longer than they are wide and have a shaft and two ends. The diaphysis, or central shaft, contains bone marrow in a marrow cavity. The rounded ends, the epiphyses, are covered with articular cartilage and are filled with red bone marrow, which produces blood cells (Figure \(2\)). Most of the limb bones are long bones—for example, the femur, tibia, ulna, and radius. Exceptions to this include the patella and the bones of the wrist and ankle.
Short bones, or cuboidal bones, are bones that are the same width and length, giving them a cube-like shape. For example, the bones of the wrist (carpals) and ankle (tarsals) are short bones (Figure \(1\)).
Flat bones are thin and relatively broad bones that are found where extensive protection of organs is required or where broad surfaces of muscle attachment are required. Examples of flat bones are the sternum (breast bone), ribs, scapulae (shoulder blades), and the roof of the skull (Figure \(1\)).
Irregular bones are bones with complex shapes. These bones may have short, flat, notched, or ridged surfaces. Examples of irregular bones are the vertebrae, hip bones, and several skull bones.
Sesamoid bones are small, flat bones and are shaped similarly to a sesame seed. The patellae are sesamoid bones (Figure \(3\)). Sesamoid bones develop inside tendons and may be found near joints at the knees, hands, and feet.
Sutural bones are small, flat, irregularly shaped bones. They may be found between the flat bones of the skull. They vary in number, shape, size, and position.
Bone Tissue
Bones are considered organs because they contain various types of tissue, such as blood, connective tissue, nerves, and bone tissue. Osteocytes, the living cells of bone tissue, form the mineral matrix of bones. There are two types of bone tissue: compact and spongy.
Compact Bone Tissue
Compact bone (or cortical bone) forms the hard external layer of all bones and surrounds the medullary cavity, or bone marrow. It provides protection and strength to bones. Compact bone tissue consists of units called osteons or Haversian systems. Osteons are cylindrical structures that contain a mineral matrix and living osteocytes connected by canaliculi, which transport blood. They are aligned parallel to the long axis of the bone. Each osteon consists of lamellae, which are layers of compact matrix that surround a central canal called the Haversian canal. The Haversian canal (osteonic canal) contains the bone’s blood vessels and nerve fibers (Figure \(4\)). Osteons in compact bone tissue are aligned in the same direction along lines of stress and help the bone resist bending or fracturing. Therefore, compact bone tissue is prominent in areas of bone at which stresses are applied in only a few directions.
Art Connection
Which of the following statements about bone tissue is false?
1. Compact bone tissue is made of cylindrical osteons that are aligned such that they travel the length of the bone.
2. Haversian canals contain blood vessels only.
3. Haversian canals contain blood vessels and nerve fibers.
4. Spongy tissue is found on the interior of the bone, and compact bone tissue is found on the exterior.
Spongy Bone Tissue
Whereas compact bone tissue forms the outer layer of all bones, spongy bone or cancellous bone forms the inner layer of all bones. Spongy bone tissue does not contain osteons that constitute compact bone tissue. Instead, it consists of trabeculae, which are lamellae that are arranged as rods or plates. Red bone marrow is found between the trabuculae. Blood vessels within this tissue deliver nutrients to osteocytes and remove waste. The red bone marrow of the femur and the interior of other large bones, such as the ileum, forms blood cells.
Spongy bone reduces the density of bone and allows the ends of long bones to compress as the result of stresses applied to the bone. Spongy bone is prominent in areas of bones that are not heavily stressed or where stresses arrive from many directions. The epiphyses of bones, such as the neck of the femur, are subject to stress from many directions. Imagine laying a heavy framed picture flat on the floor. You could hold up one side of the picture with a toothpick if the toothpick was perpendicular to the floor and the picture. Now drill a hole and stick the toothpick into the wall to hang up the picture. In this case, the function of the toothpick is to transmit the downward pressure of the picture to the wall. The force on the picture is straight down to the floor, but the force on the toothpick is both the picture wire pulling down and the bottom of the hole in the wall pushing up. The toothpick will break off right at the wall.
The neck of the femur is horizontal like the toothpick in the wall. The weight of the body pushes it down near the joint, but the vertical diaphysis of the femur pushes it up at the other end. The neck of the femur must be strong enough to transfer the downward force of the body weight horizontally to the vertical shaft of the femur (Figure \(5\)).
Link to Learning
View micrographs of musculoskeletal tissues as you review the anatomy.
Cell Types in Bones
Bone consists of four types of cells: osteoblasts, osteoclasts, osteocytes, and osteoprogenitor cells. Osteoblasts are bone cells that are responsible for bone formation. Osteoblasts synthesize and secrete the organic part and inorganic part of the extracellular matrix of bone tissue, and collagen fibers. Osteoblasts become trapped in these secretions and differentiate into less active osteocytes. Osteoclasts are large bone cells with up to 50 nuclei. They remove bone structure by releasing lysosomal enzymes and acids that dissolve the bony matrix. These minerals, released from bones into the blood, help regulate calcium concentrations in body fluids. Bone may also be resorbed for remodeling, if the applied stresses have changed. Osteocytes are mature bone cells and are the main cells in bony connective tissue; these cells cannot divide. Osteocytes maintain normal bone structure by recycling the mineral salts in the bony matrix. Osteoprogenitor cells are squamous stem cells that divide to produce daughter cells that differentiate into osteoblasts. Osteoprogenitor cells are important in the repair of fractures.
Development of Bone
Ossification, or osteogenesis, is the process of bone formation by osteoblasts. Ossification is distinct from the process of calcification; whereas calcification takes place during the ossification of bones, it can also occur in other tissues. Ossification begins approximately six weeks after fertilization in an embryo. Before this time, the embryonic skeleton consists entirely of fibrous membranes and hyaline cartilage. The development of bone from fibrous membranes is called intramembranous ossification; development from hyaline cartilage is called endochondral ossification. Bone growth continues until approximately age 25. Bones can grow in thickness throughout life, but after age 25, ossification functions primarily in bone remodeling and repair.
Intramembranous Ossification
Intramembranous ossification is the process of bone development from fibrous membranes. It is involved in the formation of the flat bones of the skull, the mandible, and the clavicles. Ossification begins as mesenchymal cells form a template of the future bone. They then differentiate into osteoblasts at the ossification center. Osteoblasts secrete the extracellular matrix and deposit calcium, which hardens the matrix. The non-mineralized portion of the bone or osteoid continues to form around blood vessels, forming spongy bone. Connective tissue in the matrix differentiates into red bone marrow in the fetus. The spongy bone is remodeled into a thin layer of compact bone on the surface of the spongy bone.
Endochondral Ossification
Endochondral ossification is the process of bone development from hyaline cartilage. All of the bones of the body, except for the flat bones of the skull, mandible, and clavicles, are formed through endochondral ossification.
In long bones, chondrocytes form a template of the hyaline cartilage diaphysis. Responding to complex developmental signals, the matrix begins to calcify. This calcification prevents diffusion of nutrients into the matrix, resulting in chondrocytes dying and the opening up of cavities in the diaphysis cartilage. Blood vessels invade the cavities, and osteoblasts and osteoclasts modify the calcified cartilage matrix into spongy bone. Osteoclasts then break down some of the spongy bone to create a marrow, or medullary, cavity in the center of the diaphysis. Dense, irregular connective tissue forms a sheath (periosteum) around the bones. The periosteum assists in attaching the bone to surrounding tissues, tendons, and ligaments. The bone continues to grow and elongate as the cartilage cells at the epiphyses divide.
In the last stage of prenatal bone development, the centers of the epiphyses begin to calcify. Secondary ossification centers form in the epiphyses as blood vessels and osteoblasts enter these areas and convert hyaline cartilage into spongy bone. Until adolescence, hyaline cartilage persists at the epiphyseal plate (growth plate), which is the region between the diaphysis and epiphysis that is responsible for the lengthwise growth of long bones (Figure \(6\)).
Growth of Bone
Long bones continue to lengthen, potentially until adolescence, through the addition of bone tissue at the epiphyseal plate. They also increase in width through appositional growth.
Lengthening of Long Bones
Chondrocytes on the epiphyseal side of the epiphyseal plate divide; one cell remains undifferentiated near the epiphysis, and one cell moves toward the diaphysis. The cells, which are pushed from the epiphysis, mature and are destroyed by calcification. This process replaces cartilage with bone on the diaphyseal side of the plate, resulting in a lengthening of the bone.
Long bones stop growing at around the age of 18 in females and the age of 21 in males in a process called epiphyseal plate closure. During this process, cartilage cells stop dividing and all of the cartilage is replaced by bone. The epiphyseal plate fades, leaving a structure called the epiphyseal line or epiphyseal remnant, and the epiphysis and diaphysis fuse.
Thickening of Long Bones
Appositional growth is the increase in the diameter of bones by the addition of bony tissue at the surface of bones. Osteoblasts at the bone surface secrete bone matrix, and osteoclasts on the inner surface break down bone. The osteoblasts differentiate into osteocytes. A balance between these two processes allows the bone to thicken without becoming too heavy.
Bone Remodeling and Repair
Bone renewal continues after birth into adulthood. Bone remodeling is the replacement of old bone tissue by new bone tissue. It involves the processes of bone deposition by osteoblasts and bone resorption by osteoclasts. Normal bone growth requires vitamins D, C, and A, plus minerals such as calcium, phosphorous, and magnesium. Hormones such as parathyroid hormone, growth hormone, and calcitonin are also required for proper bone growth and maintenance.
Bone turnover rates are quite high, with five to seven percent of bone mass being recycled every week. Differences in turnover rate exist in different areas of the skeleton and in different areas of a bone. For example, the bone in the head of the femur may be fully replaced every six months, whereas the bone along the shaft is altered much more slowly.
Bone remodeling allows bones to adapt to stresses by becoming thicker and stronger when subjected to stress. Bones that are not subject to normal stress, for example when a limb is in a cast, will begin to lose mass. A fractured or broken bone undergoes repair through four stages:
1. Blood vessels in the broken bone tear and hemorrhage, resulting in the formation of clotted blood, or a hematoma, at the site of the break. The severed blood vessels at the broken ends of the bone are sealed by the clotting process, and bone cells that are deprived of nutrients begin to die.
2. Within days of the fracture, capillaries grow into the hematoma, and phagocytic cells begin to clear away the dead cells. Though fragments of the blood clot may remain, fibroblasts and osteoblasts enter the area and begin to reform bone. Fibroblasts produce collagen fibers that connect the broken bone ends, and osteoblasts start to form spongy bone. The repair tissue between the broken bone ends is called the fibrocartilaginous callus, as it is composed of both hyaline and fibrocartilage (Figure \(7\)). Some bone spicules may also appear at this point.
3. The fibrocartilaginous callus is converted into a bony callus of spongy bone. It takes about two months for the broken bone ends to be firmly joined together after the fracture. This is similar to the endochondral formation of bone, as cartilage becomes ossified; osteoblasts, osteoclasts, and bone matrix are present.
4. The bony callus is then remodelled by osteoclasts and osteoblasts, with excess material on the exterior of the bone and within the medullary cavity being removed. Compact bone is added to create bone tissue that is similar to the original, unbroken bone. This remodeling can take many months, and the bone may remain uneven for years.
Scientific Method Connection: Decalcification of Bones
Question: What effect does the removal of calcium and collagen have on bone structure?
Background: Conduct a literature search on the role of calcium and collagen in maintaining bone structure. Conduct a literature search on diseases in which bone structure is compromised.
Hypothesis: Develop a hypothesis that states predictions of the flexibility, strength, and mass of bones that have had the calcium and collagen components removed. Develop a hypothesis regarding the attempt to add calcium back to decalcified bones.
Test the hypothesis: Test the prediction by removing calcium from chicken bones by placing them in a jar of vinegar for seven days. Test the hypothesis regarding adding calcium back to decalcified bone by placing the decalcified chicken bones into a jar of water with calcium supplements added. Test the prediction by denaturing the collagen from the bones by baking them at 250°C for three hours.
Analyze the data: Create a table showing the changes in bone flexibility, strength, and mass in the three different environments.
Report the results: Under which conditions was the bone most flexible? Under which conditions was the bone the strongest?
Draw a conclusion: Did the results support or refute the hypothesis? How do the results observed in this experiment correspond to diseases that destroy bone tissue?
Summary
Bone, or osseous tissue, is connective tissue that includes specialized cells, mineral salts, and collagen fibers. The human skeleton can be divided into long bones, short bones, flat bones, and irregular bones. Compact bone tissue is composed of osteons and forms the external layer of all bones. Spongy bone tissue is composed of trabeculae and forms the inner part of all bones. Four types of cells compose bony tissue: osteocytes, osteoclasts, osteoprogenitor cells, and osteoblasts. Ossification is the process of bone formation by osteoblasts. Intramembranous ossification is the process of bone development from fibrous membranes. Endochondral ossification is the process of bone development from hyaline cartilage. Long bones lengthen as chondrocytes divide and secrete hyaline cartilage. Osteoblasts replace cartilage with bone. Appositional growth is the increase in the diameter of bones by the addition of bone tissue at the surface of bones. Bone remodeling involves the processes of bone deposition by osteoblasts and bone resorption by osteoclasts. Bone repair occurs in four stages and can take several months.
Art Exercise
Figure \(4\): Which of the following statements about bone tissue is false?
1. Compact bone tissue is made of cylindrical osteons that are aligned such that they travel the length of the bone.
2. Haversian canals contain blood vessels only.
3. Haversian canals contain blood vessels and nerve fibers.
4. Spongy tissue is found on the interior of the bone, and compact bone tissue is found on the exterior.
Answer
B
Glossary
appositional growth
increase in the diameter of bones by the addition of bone tissue at the surface of bones
bone
(also, osseous tissue) connective tissue that constitutes the endoskeleton
bone remodeling
replacement of old bone tissue by new bone tissue
calcification
process of deposition of mineral salts in the collagen fiber matrix that crystallizes and hardens the tissue
compact bone
forms the hard external layer of all bones
diaphysis
central shaft of bone, contains bone marrow in a marrow cavity
endochondral ossification
process of bone development from hyaline cartilage
epiphyseal plate
region between the diaphysis and epiphysis that is responsible for the lengthwise growth of long bones
epiphysis
rounded end of bone, covered with articular cartilage and filled with red bone marrow, which produces blood cells
flat bone
thin and relatively broad bone found where extensive protection of organs is required or where broad surfaces of muscle attachment are required
Haversian canal
contains the bone’s blood vessels and nerve fibers
intramembranous ossification
process of bone development from fibrous membranes
irregular bone
bone with complex shapes; examples include vertebrae and hip bones
lamella
layer of compact tissue that surrounds a central canal called the Haversian canal
long bone
bone that is longer than wide, and has a shaft and two ends
osteoblast
bone cell responsible for bone formation
osteoclast
large bone cells with up to 50 nuclei, responsible for bone remodeling
osteocyte
mature bone cells and the main cell in bone tissue
osseous tissue
connective tissue that constitutes the endoskeleton
ossification
(also, osteogenesis) process of bone formation by osteoblasts
osteon
cylindrical structure aligned parallel to the long axis of the bone
resorption
process by which osteoclasts release minerals stored in bones
sesamoid bone
small, flat bone shaped like a sesame seed; develops inside tendons
short bone
bone that has the same width and length, giving it a cube-like shape
spongy bone tissue
forms the inner layer of all bones
suture bone
small, flat, irregularly shaped bone that forms between the flat bones of the cranium
trabeculae
lamellae that are arranged as rods or plates | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/45%3A_The_Musculoskeletal_System/45.02%3A_A_Closer_Look_at_Bone.txt |
Skills to Develop
• Classify the different types of joints on the basis of structure
• Explain the role of joints in skeletal movement
The point at which two or more bones meet is called a joint, or articulation. Joints are responsible for movement, such as the movement of limbs, and stability, such as the stability found in the bones of the skull.
Classification of Joints on the Basis of Structure
There are two ways to classify joints: on the basis of their structure or on the basis of their function. The structural classification divides joints into bony, fibrous, cartilaginous, and synovial joints depending on the material composing the joint and the presence or absence of a cavity in the joint.
Fibrous Joints
The bones of fibrous joints are held together by fibrous connective tissue. There is no cavity, or space, present between the bones and so most fibrous joints do not move at all, or are only capable of minor movements. There are three types of fibrous joints: sutures, syndesmoses, and gomphoses. Sutures are found only in the skull and possess short fibers of connective tissue that hold the skull bones tightly in place (Figure \(1\)).
Syndesmoses are joints in which the bones are connected by a band of connective tissue, allowing for more movement than in a suture. An example of a syndesmosis is the joint of the tibia and fibula in the ankle. The amount of movement in these types of joints is determined by the length of the connective tissue fibers. Gomphoses occur between teeth and their sockets; the term refers to the way the tooth fits into the socket like a peg (Figure \(2\)). The tooth is connected to the socket by a connective tissue referred to as the periodontal ligament.
Cartilaginous Joints
Cartilaginous joints are joints in which the bones are connected by cartilage. There are two types of cartilaginous joints: synchondroses and symphyses. In a synchondrosis, the bones are joined by hyaline cartilage. Synchondroses are found in the epiphyseal plates of growing bones in children. In symphyses, hyaline cartilage covers the end of the bone but the connection between bones occurs through fibrocartilage. Symphyses are found at the joints between vertebrae. Either type of cartilaginous joint allows for very little movement.
Synovial Joints
Synovial joints are the only joints that have a space between the adjoining bones (Figure \(3\)). This space is referred to as the synovial (or joint) cavity and is filled with synovial fluid. Synovial fluid lubricates the joint, reducing friction between the bones and allowing for greater movement. The ends of the bones are covered with articular cartilage, a hyaline cartilage, and the entire joint is surrounded by an articular capsule composed of connective tissue that allows movement of the joint while resisting dislocation. Articular capsules may also possess ligaments that hold the bones together. Synovial joints are capable of the greatest movement of the three structural joint types; however, the more mobile a joint, the weaker the joint. Knees, elbows, and shoulders are examples of synovial joints.
Classification of Joints on the Basis of Function
The functional classification divides joints into three categories: synarthroses, amphiarthroses, and diarthroses. A synarthrosis is a joint that is immovable. This includes sutures, gomphoses, and synchondroses. Amphiarthroses are joints that allow slight movement, including syndesmoses and symphyses. Diarthroses are joints that allow for free movement of the joint, as in synovial joints.
Movement at Synovial Joints
The wide range of movement allowed by synovial joints produces different types of movements. The movement of synovial joints can be classified as one of four different types: gliding, angular, rotational, or special movement.
Gliding Movement
Gliding movements occur as relatively flat bone surfaces move past each other. Gliding movements produce very little rotation or angular movement of the bones. The joints of the carpal and tarsal bones are examples of joints that produce gliding movements.
Angular Movement
Angular movements are produced when the angle between the bones of a joint changes. There are several different types of angular movements, including flexion, extension, hyperextension, abduction, adduction, and circumduction. Flexion, or bending, occurs when the angle between the bones decreases. Moving the forearm upward at the elbow or moving the wrist to move the hand toward the forearm are examples of flexion. Extension is the opposite of flexion in that the angle between the bones of a joint increases. Straightening a limb after flexion is an example of extension. Extension past the regular anatomical position is referred to as hyperextension. This includes moving the neck back to look upward, or bending the wrist so that the hand moves away from the forearm.
Abduction occurs when a bone moves away from the midline of the body. Examples of abduction are moving the arms or legs laterally to lift them straight out to the side. Adduction is the movement of a bone toward the midline of the body. Movement of the limbs inward after abduction is an example of adduction. Circumduction is the movement of a limb in a circular motion, as in moving the arm in a circular motion.
Rotational Movement
Rotational movement is the movement of a bone as it rotates around its longitudinal axis. Rotation can be toward the midline of the body, which is referred to as medial rotation, or away from the midline of the body, which is referred to as lateral rotation. Movement of the head from side to side is an example of rotation.
Special Movements
Some movements that cannot be classified as gliding, angular, or rotational are called special movements. Inversion involves the soles of the feet moving inward, toward the midline of the body. Eversion is the opposite of inversion, movement of the sole of the foot outward, away from the midline of the body. Protraction is the anterior movement of a bone in the horizontal plane. Retraction occurs as a joint moves back into position after protraction. Protraction and retraction can be seen in the movement of the mandible as the jaw is thrust outwards and then back inwards. Elevation is the movement of a bone upward, such as when the shoulders are shrugged, lifting the scapulae. Depression is the opposite of elevation—movement downward of a bone, such as after the shoulders are shrugged and the scapulae return to their normal position from an elevated position. Dorsiflexion is a bending at the ankle such that the toes are lifted toward the knee. Plantar flexion is a bending at the ankle when the heel is lifted, such as when standing on the toes. Supination is the movement of the radius and ulna bones of the forearm so that the palm faces forward. Pronation is the opposite movement, in which the palm faces backward. Opposition is the movement of the thumb toward the fingers of the same hand, making it possible to grasp and hold objects.
Types of Synovial Joints
Synovial joints are further classified into six different categories on the basis of the shape and structure of the joint. The shape of the joint affects the type of movement permitted by the joint (Figure \(4\)). These joints can be described as planar, hinge, pivot, condyloid, saddle, or ball-and-socket joints.
Planar Joints
Planar joints have bones with articulating surfaces that are flat or slightly curved faces. These joints allow for gliding movements, and so the joints are sometimes referred to as gliding joints. The range of motion is limited in these joints and does not involve rotation. Planar joints are found in the carpal bones in the hand and the tarsal bones of the foot, as well as between vertebrae (Figure \(5\)).
Hinge Joints
In hinge joints, the slightly rounded end of one bone fits into the slightly hollow end of the other bone. In this way, one bone moves while the other remains stationary, like the hinge of a door. The elbow is an example of a hinge joint. The knee is sometimes classified as a modified hinge joint (Figure \(6\)).
Pivot Joints
Pivot joints consist of the rounded end of one bone fitting into a ring formed by the other bone. This structure allows rotational movement, as the rounded bone moves around its own axis. An example of a pivot joint is the joint of the first and second vertebrae of the neck that allows the head to move back and forth (Figure \(7\)). The joint of the wrist that allows the palm of the hand to be turned up and down is also a pivot joint.
Condyloid Joints
Condyloid joints consist of an oval-shaped end of one bone fitting into a similarly oval-shaped hollow of another bone (Figure \(8\)). This is also sometimes called an ellipsoidal joint. This type of joint allows angular movement along two axes, as seen in the joints of the wrist and fingers, which can move both side to side and up and down.
Saddle Joints
Saddle joints are so named because the ends of each bone resemble a saddle, with concave and convex portions that fit together. Saddle joints allow angular movements similar to condyloid joints but with a greater range of motion. An example of a saddle joint is the thumb joint, which can move back and forth and up and down, but more freely than the wrist or fingers (Figure \(9\)).
Ball-and-Socket Joints
Ball-and-socket joints possess a rounded, ball-like end of one bone fitting into a cuplike socket of another bone. This organization allows the greatest range of motion, as all movement types are possible in all directions. Examples of ball-and-socket joints are the shoulder and hip joints (Figure \(10\)).
Link to Learning
Watch this animation showing the six types of synovial joints.
Career Connection: Rheumatologist
Rheumatologists are medical doctors who specialize in the diagnosis and treatment of disorders of the joints, muscles, and bones. They diagnose and treat diseases such as arthritis, musculoskeletal disorders, osteoporosis, and autoimmune diseases such as ankylosing spondylitis and rheumatoid arthritis.
Rheumatoid arthritis (RA) is an inflammatory disorder that primarily affects the synovial joints of the hands, feet, and cervical spine. Affected joints become swollen, stiff, and painful. Although it is known that RA is an autoimmune disease in which the body’s immune system mistakenly attacks healthy tissue, the cause of RA remains unknown. Immune cells from the blood enter joints and the synovium causing cartilage breakdown, swelling, and inflammation of the joint lining. Breakdown of cartilage causes bones to rub against each other causing pain. RA is more common in women than men and the age of onset is usually 40–50 years of age.
Rheumatologists can diagnose RA on the basis of symptoms such as joint inflammation and pain, X-ray and MRI imaging, and blood tests. Arthrography is a type of medical imaging of joints that uses a contrast agent, such as a dye, that is opaque to X-rays. This allows the soft tissue structures of joints—such as cartilage, tendons, and ligaments—to be visualized. An arthrogram differs from a regular X-ray by showing the surface of soft tissues lining the joint in addition to joint bones. An arthrogram allows early degenerative changes in joint cartilage to be detected before bones become affected.
There is currently no cure for RA; however, rheumatologists have a number of treatment options available. Early stages can be treated with rest of the affected joints by using a cane or by using joint splints that minimize inflammation. When inflammation has decreased, exercise can be used to strengthen the muscles that surround the joint and to maintain joint flexibility. If joint damage is more extensive, medications can be used to relieve pain and decrease inflammation. Anti-inflammatory drugs such as aspirin, topical pain relievers, and corticosteroid injections may be used. Surgery may be required in cases in which joint damage is severe.
Summary
The structural classification of joints divides them into bony, fibrous, cartilaginous, and synovial joints. The bones of fibrous joints are held together by fibrous connective tissue; the three types of fibrous joints are sutures, syndesomes, and gomphoses. Cartilaginous joints are joints in which the bones are connected by cartilage; the two types of cartilaginous joints are synchondroses and symphyses. Synovial joints are joints that have a space between the adjoining bones. The functional classification divides joints into three categories: synarthroses, amphiarthroses, and diarthroses. The movement of synovial joints can be classified as one of four different types: gliding, angular, rotational, or special movement. Gliding movements occur as relatively flat bone surfaces move past each other. Angular movements are produced when the angle between the bones of a joint changes. Rotational movement is the movement of a bone as it rotates around its own longitudinal axis. Special movements include inversion, eversion, protraction, retraction, elevation, depression, dorsiflexion, plantar flexion, supination, pronation, and opposition. Synovial joints are also classified into six different categories on the basis of the shape and structure of the joint: planar, hinge, pivot, condyloid, saddle, and ball-and-socket.
Glossary
abduction
when a bone moves away from the midline of the body
adduction
movement of the limbs inward after abduction
amphiarthrosis
joint that allows slight movement; includes syndesmoses and symphyses
angular movement
produced when the angle between the bones of a joint changes
ball-and-socket joint
joint with a rounded, ball-like end of one bone fitting into a cuplike socket of another bone
cartilaginous joint
joint in which the bones are connected by cartilage
circumduction
movement of a limb in a circular motion.
condyloid joint
oval-shaped end of one bone fitting into a similarly oval-shaped hollow of another bone
depression
movement downward of a bone, such as after the shoulders are shrugged and the scapulae return to their normal position from an elevated position; opposite of elevation
diarthrosis
joint that allows for free movement of the joint; found in synovial joints
dorsiflexion
bending at the ankle such that the toes are lifted toward the knee
elevation
movement of a bone upward, such as when the shoulders are shrugged, lifting the scapulae
eversion
movement of the sole of the foot outward, away from the midline of the body; opposite of inversion
extension
movement in which the angle between the bones of a joint increases; opposite of flexion
fibrous joint
joint held together by fibrous connective tissue
flexion
movement in which the angle between the bones decreases; opposite of extension
gliding movement
when relatively flat bone surfaces move past each other
gomphosis
the joint in which the tooth fits into the socket like a peg
hinge joint
slightly rounded end of one bone fits into the slightly hollow end of the other bone
hyperextension
extension past the regular anatomical position
inversion
soles of the feet moving inward, toward the midline of the body
joint
point at which two or more bones meet
lateral rotation
rotation away from the midline of the body
medial rotation
rotation toward the midline of the body
opposition
movement of the thumb toward the fingers of the same hand, making it possible to grasp and hold objects
plantar flexion
bending at the ankle such that the heel is lifted, such as when standing on the toes
planar joint
joint with bones whose articulating surfaces are flat
pivot joint
joint with the rounded end of one bone fitting into a ring formed by the other bone
pronation
movement in which the palm faces backward
protraction
anterior movement of a bone in the horizontal plane
retraction
movement in which a joint moves back into position after protraction
rotational movement
movement of a bone as it rotates around its own longitudinal axis
saddle joint
joint with concave and convex portions that fit together; named because the ends of each bone resemble a saddle
supination
movement of the radius and ulna bones of the forearm so that the palm faces forward
suture
short fiber of connective tissue that holds the skull bones tightly in place; found only in the skull
synarthrosis
joint that is immovable
symphysis
hyaline cartilage covers the end of the bone, but the connection between bones occurs through fibrocartilage; symphyses are found at the joints between vertebrae
synchondrosis
bones joined by hyaline cartilage; synchondroses are found in the epiphyseal plates of growing bones in children
syndesmosis
joint in which the bones are connected by a band of connective tissue, allowing for more movement than in a suture
synovial joint
only joint that has a space between the adjoining bones | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/45%3A_The_Musculoskeletal_System/45.03%3A_Joints.txt |
Skills to Develop
• Classify the different types of muscle tissue
• Explain the role of muscles in locomotion
Muscle cells are specialized for contraction. Muscles allow for motions such as walking, and they also facilitate bodily processes such as respiration and digestion. The body contains three types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle (Figure \(1\)).
Skeletal muscle tissue forms skeletal muscles, which attach to bones or skin and control locomotion and any movement that can be consciously controlled. Because it can be controlled by thought, skeletal muscle is also called voluntary muscle. Skeletal muscles are long and cylindrical in appearance; when viewed under a microscope, skeletal muscle tissue has a striped or striated appearance. The striations are caused by the regular arrangement of contractile proteins (actin and myosin). Actin is a globular contractile protein that interacts with myosin for muscle contraction. Skeletal muscle also has multiple nuclei present in a single cell.
Smooth muscle tissue occurs in the walls of hollow organs such as the intestines, stomach, and urinary bladder, and around passages such as the respiratory tract and blood vessels. Smooth muscle has no striations, is not under voluntary control, has only one nucleus per cell, is tapered at both ends, and is called involuntary muscle.
Cardiac muscle tissue is only found in the heart, and cardiac contractions pump blood throughout the body and maintain blood pressure. Like skeletal muscle, cardiac muscle is striated, but unlike skeletal muscle, cardiac muscle cannot be consciously controlled and is called involuntary muscle. It has one nucleus per cell, is branched, and is distinguished by the presence of intercalated disks.
Skeletal Muscle Fiber Structure
Each skeletal muscle fiber is a skeletal muscle cell. These cells are incredibly large, with diameters of up to 100 µm and lengths of up to 30 cm. The plasma membrane of a skeletal muscle fiber is called the sarcolemma. The sarcolemma is the site of action potential conduction, which triggers muscle contraction. Within each muscle fiber are myofibrils—long cylindrical structures that lie parallel to the muscle fiber. Myofibrils run the entire length of the muscle fiber, and because they are only approximately 1.2 µm in diameter, hundreds to thousands can be found inside one muscle fiber. They attach to the sarcolemma at their ends, so that as myofibrils shorten, the entire muscle cell contracts (Figure \(2\)).
The striated appearance of skeletal muscle tissue is a result of repeating bands of the proteins actin and myosin that are present along the length of myofibrils. Dark A bands and light I bands repeat along myofibrils, and the alignment of myofibrils in the cell causes the entire cell to appear striated or banded.
Each I band has a dense line running vertically through the middle called a Z disc or Z line. The Z discs mark the border of units called sarcomeres, which are the functional units of skeletal muscle. One sarcomere is the space between two consecutive Z discs and contains one entire A band and two halves of an I band, one on either side of the A band. A myofibril is composed of many sarcomeres running along its length, and as the sarcomeres individually contract, the myofibrils and muscle cells shorten (Figure \(3\)).
Myofibrils are composed of smaller structures called myofilaments. There are two main types of filaments: thick filaments and thin filaments; each has different compositions and locations. Thick filaments occur only in the A band of a myofibril. Thin filaments attach to a protein in the Z disc called alpha-actinin and occur across the entire length of the I band and partway into the A band. The region at which thick and thin filaments overlap has a dense appearance, as there is little space between the filaments. Thin filaments do not extend all the way into the A bands, leaving a central region of the A band that only contains thick filaments. This central region of the A band looks slightly lighter than the rest of the A band and is called the H zone. The middle of the H zone has a vertical line called the M line, at which accessory proteins hold together thick filaments. Both the Z disc and the M line hold myofilaments in place to maintain the structural arrangement and layering of the myofibril. Myofibrils are connected to each other by intermediate, or desmin, filaments that attach to the Z disc.
Thick and thin filaments are themselves composed of proteins. Thick filaments are composed of the protein myosin. The tail of a myosin molecule connects with other myosin molecules to form the central region of a thick filament near the M line, whereas the heads align on either side of the thick filament where the thin filaments overlap. The primary component of thin filaments is the actin protein. Two other components of the thin filament are tropomyosin and troponin. Actin has binding sites for myosin attachment. Strands of tropomyosin block the binding sites and prevent actin–myosin interactions when the muscles are at rest. Troponin consists of three globular subunits. One subunit binds to tropomyosin, one subunit binds to actin, and one subunit binds Ca2+ ions.
Link to Learning
View this animation showing the organization of muscle fibers.
Sliding Filament Model of Contraction
For a muscle cell to contract, the sarcomere must shorten. However, thick and thin filaments—the components of sarcomeres—do not shorten. Instead, they slide by one another, causing the sarcomere to shorten while the filaments remain the same length. The sliding filament theory of muscle contraction was developed to fit the differences observed in the named bands on the sarcomere at different degrees of muscle contraction and relaxation. The mechanism of contraction is the binding of myosin to actin, forming cross-bridges that generate filament movement (Figure \(4\)).
When a sarcomere shortens, some regions shorten whereas others stay the same length. A sarcomere is defined as the distance between two consecutive Z discs or Z lines; when a muscle contracts, the distance between the Z discs is reduced. The H zone—the central region of the A zone—contains only thick filaments and is shortened during contraction. The I band contains only thin filaments and also shortens. The A band does not shorten—it remains the same length—but A bands of different sarcomeres move closer together during contraction, eventually disappearing. Thin filaments are pulled by the thick filaments toward the center of the sarcomere until the Z discs approach the thick filaments. The zone of overlap, in which thin filaments and thick filaments occupy the same area, increases as the thin filaments move inward.
ATP and Muscle Contraction
The motion of muscle shortening occurs as myosin heads bind to actin and pull the actin inwards. This action requires energy, which is provided by ATP. Myosin binds to actin at a binding site on the globular actin protein. Myosin has another binding site for ATP at which enzymatic activity hydrolyzes ATP to ADP, releasing an inorganic phosphate molecule and energy.
ATP binding causes myosin to release actin, allowing actin and myosin to detach from each other. After this happens, the newly bound ATP is converted to ADP and inorganic phosphate, Pi. The enzyme at the binding site on myosin is called ATPase. The energy released during ATP hydrolysis changes the angle of the myosin head into a “cocked” position. The myosin head is then in a position for further movement, possessing potential energy, but ADP and Pi are still attached. If actin binding sites are covered and unavailable, the myosin will remain in the high energy configuration with ATP hydrolyzed, but still attached.
If the actin binding sites are uncovered, a cross-bridge will form; that is, the myosin head spans the distance between the actin and myosin molecules. Pi is then released, allowing myosin to expend the stored energy as a conformational change. The myosin head moves toward the M line, pulling the actin along with it. As the actin is pulled, the filaments move approximately 10 nm toward the M line. This movement is called the power stroke, as it is the step at which force is produced. As the actin is pulled toward the M line, the sarcomere shortens and the muscle contracts.
When the myosin head is “cocked,” it contains energy and is in a high-energy configuration. This energy is expended as the myosin head moves through the power stroke; at the end of the power stroke, the myosin head is in a low-energy position. After the power stroke, ADP is released; however, the cross-bridge formed is still in place, and actin and myosin are bound together. ATP can then attach to myosin, which allows the cross-bridge cycle to start again and further muscle contraction can occur (Figure \(5\)).
Link to Learning
Watch this video explaining how a muscle contraction is signaled.
Art Connection
Which of the following statements about muscle contraction is true?
1. The power stroke occurs when ATP is hydrolyzed to ADP and phosphate.
2. The power stroke occurs when ADP and phosphate dissociate from the myosin head.
3. The power stroke occurs when ADP and phosphate dissociate from the actin active site.
4. The power stroke occurs when Ca2+ binds the calcium head.
Link to Learning
View this animation of the cross-bridge muscle contraction.
Regulatory Proteins
When a muscle is in a resting state, actin and myosin are separated. To keep actin from binding to the active site on myosin, regulatory proteins block the molecular binding sites. Tropomyosin blocks myosin binding sites on actin molecules, preventing cross-bridge formation and preventing contraction in a muscle without nervous input. Troponin binds to tropomyosin and helps to position it on the actin molecule; it also binds calcium ions.
To enable a muscle contraction, tropomyosin must change conformation, uncovering the myosin-binding site on an actin molecule and allowing cross-bridge formation. This can only happen in the presence of calcium, which is kept at extremely low concentrations in the sarcoplasm. If present, calcium ions bind to troponin, causing conformational changes in troponin that allow tropomyosin to move away from the myosin binding sites on actin. Once the tropomyosin is removed, a cross-bridge can form between actin and myosin, triggering contraction. Cross-bridge cycling continues until Ca2+ ions and ATP are no longer available and tropomyosin again covers the binding sites on actin.
Excitation–Contraction Coupling
Excitation–contraction coupling is the link (transduction) between the action potential generated in the sarcolemma and the start of a muscle contraction. The trigger for calcium release from the sarcoplasmic reticulum into the sarcoplasm is a neural signal. Each skeletal muscle fiber is controlled by a motor neuron, which conducts signals from the brain or spinal cord to the muscle. The area of the sarcolemma on the muscle fiber that interacts with the neuron is called the motor end plate. The end of the neuron’s axon is called the synaptic terminal, and it does not actually contact the motor end plate. A small space called the synaptic cleft separates the synaptic terminal from the motor end plate. Electrical signals travel along the neuron’s axon, which branches through the muscle and connects to individual muscle fibers at a neuromuscular junction.
The ability of cells to communicate electrically requires that the cells expend energy to create an electrical gradient across their cell membranes. This charge gradient is carried by ions, which are differentially distributed across the membrane. Each ion exerts an electrical influence and a concentration influence. Just as milk will eventually mix with coffee without the need to stir, ions also distribute themselves evenly, if they are permitted to do so. In this case, they are not permitted to return to an evenly mixed state.
The sodium–potassium ATPase uses cellular energy to move K+ ions inside the cell and Na+ ions outside. This alone accumulates a small electrical charge, but a big concentration gradient. There is lots of K+ in the cell and lots of Na+ outside the cell. Potassium is able to leave the cell through K+ channels that are open 90% of the time, and it does. However, Na+ channels are rarely open, so Na+ remains outside the cell. When K+ leaves the cell, obeying its concentration gradient, that effectively leaves a negative charge behind. So at rest, there is a large concentration gradient for Na+ to enter the cell, and there is an accumulation of negative charges left behind in the cell. This is the resting membrane potential. Potential in this context means a separation of electrical charge that is capable of doing work. It is measured in volts, just like a battery. However, the transmembrane potential is considerably smaller (0.07 V); therefore, the small value is expressed as millivolts (mV) or 70 mV. Because the inside of a cell is negative compared with the outside, a minus sign signifies the excess of negative charges inside the cell, −70 mV.
If an event changes the permeability of the membrane to Na+ ions, they will enter the cell. That will change the voltage. This is an electrical event, called an action potential, that can be used as a cellular signal. Communication occurs between nerves and muscles through neurotransmitters. Neuron action potentials cause the release of neurotransmitters from the synaptic terminal into the synaptic cleft, where they can then diffuse across the synaptic cleft and bind to a receptor molecule on the motor end plate. The motor end plate possesses junctional folds—folds in the sarcolemma that create a large surface area for the neurotransmitter to bind to receptors. The receptors are actually sodium channels that open to allow the passage of Na+ into the cell when they receive neurotransmitter signal.
Acetylcholine (ACh) is a neurotransmitter released by motor neurons that binds to receptors in the motor end plate. Neurotransmitter release occurs when an action potential travels down the motor neuron’s axon, resulting in altered permeability of the synaptic terminal membrane and an influx of calcium. The Ca2+ ions allow synaptic vesicles to move to and bind with the presynaptic membrane (on the neuron), and release neurotransmitter from the vesicles into the synaptic cleft. Once released by the synaptic terminal, ACh diffuses across the synaptic cleft to the motor end plate, where it binds with ACh receptors. As a neurotransmitter binds, these ion channels open, and Na+ ions cross the membrane into the muscle cell. This reduces the voltage difference between the inside and outside of the cell, which is called depolarization. As ACh binds at the motor end plate, this depolarization is called an end-plate potential. The depolarization then spreads along the sarcolemma, creating an action potential as sodium channels adjacent to the initial depolarization site sense the change in voltage and open. The action potential moves across the entire cell, creating a wave of depolarization.
ACh is broken down by the enzyme acetylcholinesterase (AChE) into acetyl and choline. AChE resides in the synaptic cleft, breaking down ACh so that it does not remain bound to ACh receptors, which would cause unwanted extended muscle contraction (Figure \(6\)).
Art Connection
The deadly nerve gas Sarin irreversibly inhibits acetycholinesterase. What effect would Sarin have on muscle contraction?
After depolarization, the membrane returns to its resting state. This is called repolarization, during which voltage-gated sodium channels close. Potassium channels continue at 90% conductance. Because the plasma membrane sodium–potassium ATPase always transports ions, the resting state (negatively charged inside relative to the outside) is restored. The period immediately following the transmission of an impulse in a nerve or muscle, in which a neuron or muscle cell regains its ability to transmit another impulse, is called the refractory period. During the refractory period, the membrane cannot generate another action potential. . The refractory period allows the voltage-sensitive ion channels to return to their resting configurations. The sodium potassium ATPase continually moves Na+ back out of the cell and K+ back into the cell, and the K+ leaks out leaving negative charge behind. Very quickly, the membrane repolarizes, so that it can again be depolarized.
Control of Muscle Tension
Neural control initiates the formation of actin–myosin cross-bridges, leading to the sarcomere shortening involved in muscle contraction. These contractions extend from the muscle fiber through connective tissue to pull on bones, causing skeletal movement. The pull exerted by a muscle is called tension, and the amount of force created by this tension can vary. This enables the same muscles to move very light objects and very heavy objects. In individual muscle fibers, the amount of tension produced depends on the cross-sectional area of the muscle fiber and the frequency of neural stimulation.
The number of cross-bridges formed between actin and myosin determine the amount of tension that a muscle fiber can produce. Cross-bridges can only form where thick and thin filaments overlap, allowing myosin to bind to actin. If more cross-bridges are formed, more myosin will pull on actin, and more tension will be produced.
The ideal length of a sarcomere during production of maximal tension occurs when thick and thin filaments overlap to the greatest degree. If a sarcomere at rest is stretched past an ideal resting length, thick and thin filaments do not overlap to the greatest degree, and fewer cross-bridges can form. This results in fewer myosin heads pulling on actin, and less tension is produced. As a sarcomere is shortened, the zone of overlap is reduced as the thin filaments reach the H zone, which is composed of myosin tails. Because it is myosin heads that form cross-bridges, actin will not bind to myosin in this zone, reducing the tension produced by this myofiber. If the sarcomere is shortened even more, thin filaments begin to overlap with each other—reducing cross-bridge formation even further, and producing even less tension. Conversely, if the sarcomere is stretched to the point at which thick and thin filaments do not overlap at all, no cross-bridges are formed and no tension is produced. This amount of stretching does not usually occur because accessory proteins, internal sensory nerves, and connective tissue oppose extreme stretching.
The primary variable determining force production is the number of myofibers within the muscle that receive an action potential from the neuron that controls that fiber. When using the biceps to pick up a pencil, the motor cortex of the brain only signals a few neurons of the biceps, and only a few myofibers respond. In vertebrates, each myofiber responds fully if stimulated. When picking up a piano, the motor cortex signals all of the neurons in the biceps and every myofiber participates. This is close to the maximum force the muscle can produce. As mentioned above, increasing the frequency of action potentials (the number of signals per second) can increase the force a bit more, because the tropomyosin is flooded with calcium.
Summary
The body contains three types of muscle tissue: skeletal muscle, cardiac muscle, and smooth muscle. Skeleton muscle tissue is composed of sarcomeres, the functional units of muscle tissue. Muscle contraction occurs when sarcomeres shorten, as thick and thin filaments slide past each other, which is called the sliding filament model of muscle contraction. ATP provides the energy for cross-bridge formation and filament sliding. Regulatory proteins, such as troponin and tropomyosin, control cross-bridge formation. Excitation–contraction coupling transduces the electrical signal of the neuron, via acetylcholine, to an electrical signal on the muscle membrane, which initiates force production. The number of muscle fibers contracting determines how much force the whole muscle produces.
Art Connections
Figure \(5\): Which of the following statements about muscle contraction is true?
1. The power stroke occurs when ATP is hydrolyzed to ADP and phosphate.
2. The power stroke occurs when ADP and phosphate dissociate from the myosin head.
3. The power stroke occurs when ADP and phosphate dissociate from the actin active site.
4. The power stroke occurs when Ca2+ binds the calcium head.
Answer
B
Figure \(6\): The deadly nerve gas Sarin irreversibly inhibits acetycholinesterase. What effect would Sarin have on muscle contraction?
Answer
In the presence of Sarin, acetycholine is not removed from the synapse, resulting in continuous stimulation of the muscle plasma membrane. At first, muscle activity is intense and uncontrolled, but the ion gradients dissipate, so electrical signals in the T-tubules are no longer possible. The result is paralysis, leading to death by asphyxiation.
Glossary
actin
globular contractile protein that interacts with myosin for muscle contraction
acetylcholinesterase
(AChE) enzyme that breaks down ACh into acetyl and choline
cardiac muscle
tissue muscle tissue found only in the heart; cardiac contractions pump blood throughout the body and maintain blood pressure
motor end plate
sarcolemma of the muscle fiber that interacts with the neuron
myofibril
long cylindrical structures that lie parallel to the muscle fiber
myofilament
small structures that make up myofibrils
myosin
contractile protein that interacts with actin for muscle contraction
sarcolemma
plasma membrane of a skeletal muscle fiber
sarcomere
functional unit of skeletal muscle
skeletal muscle tissue
forms skeletal muscles, which attach to bones and control locomotion and any movement that can be consciously controlled
smooth muscle
tissue occurs in the walls of hollow organs such as the intestines, stomach, and urinary bladder, and around passages such as the respiratory tract and blood vessels
thick filament
a group of myosin molecules
thin filament
two polymers of actin wound together along with tropomyosin and troponin
tropomyosin
acts to block myosin binding sites on actin molecules, preventing cross-bridge formation and preventing contraction until a muscle receives a neuron signal
troponin
binds to tropomyosin and helps to position it on the actin molecule, and also binds calcium ions | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/45%3A_The_Musculoskeletal_System/45.04%3A_Muscle_Contraction.txt |
Skills to Develop
• Discuss the different types of skeletal systems
• Explain the role of the human skeletal system
• Compare and contrast different skeletal systems
A skeletal system is necessary to support the body, protect internal organs, and allow for the movement of an organism. There are three different skeleton designs that fulfill these functions: hydrostatic skeleton, exoskeleton, and endoskeleton.
Hydrostatic Skeleton
A hydrostatic skeleton is a skeleton formed by a fluid-filled compartment within the body, called the coelom. The organs of the coelom are supported by the aqueous fluid, which also resists external compression. This compartment is under hydrostatic pressure because of the fluid and supports the other organs of the organism. This type of skeletal system is found in soft-bodied animals such as sea anemones, earthworms, Cnidaria, and other invertebrates (Figure \(1\)).
Movement in a hydrostatic skeleton is provided by muscles that surround the coelom. The muscles in a hydrostatic skeleton contract to change the shape of the coelom; the pressure of the fluid in the coelom produces movement. For example, earthworms move by waves of muscular contractions of the skeletal muscle of the body wall hydrostatic skeleton, called peristalsis, which alternately shorten and lengthen the body. Lengthening the body extends the anterior end of the organism. Most organisms have a mechanism to fix themselves in the substrate. Shortening the muscles then draws the posterior portion of the body forward. Although a hydrostatic skeleton is well-suited to invertebrate organisms such as earthworms and some aquatic organisms, it is not an efficient skeleton for terrestrial animals.
Exoskeleton
An exoskeleton is an external skeleton that consists of a hard encasement on the surface of an organism. For example, the shells of crabs and insects are exoskeletons (Figure \(2\)). This skeleton type provides defence against predators, supports the body, and allows for movement through the contraction of attached muscles. As with vertebrates, muscles must cross a joint inside the exoskeleton. Shortening of the muscle changes the relationship of the two segments of the exoskeleton. Arthropods such as crabs and lobsters have exoskeletons that consist of 30–50 percent chitin, a polysaccharide derivative of glucose that is a strong but flexible material. Chitin is secreted by the epidermal cells. The exoskeleton is further strengthened by the addition of calcium carbonate in organisms such as the lobster. Because the exoskeleton is acellular, arthropods must periodically shed their exoskeletons because the exoskeleton does not grow as the organism grows.
Endoskeleton
An endoskeleton is a skeleton that consists of hard, mineralized structures located within the soft tissue of organisms. An example of a primitive endoskeletal structure is the spicules of sponges. The bones of vertebrates are composed of tissues, whereas sponges have no true tissues (Figure \(1\)). Endoskeletons provide support for the body, protect internal organs, and allow for movement through contraction of muscles attached to the skeleton.
The human skeleton is an endoskeleton that consists of 206 bones in the adult. It has five main functions: providing support to the body, storing minerals and lipids, producing blood cells, protecting internal organs, and allowing for movement. The skeletal system in vertebrates is divided into the axial skeleton (which consists of the skull, vertebral column, and rib cage), and the appendicular skeleton (which consists of the shoulders, limb bones, the pectoral girdle, and the pelvic girdle).
Human Axial Skeleton
The axial skeleton forms the central axis of the body and includes the bones of the skull, ossicles of the middle ear, hyoid bone of the throat, vertebral column, and the thoracic cage (ribcage) (Figure \(1\)). The function of the axial skeleton is to provide support and protection for the brain, the spinal cord, and the organs in the ventral body cavity. It provides a surface for the attachment of muscles that move the head, neck, and trunk, performs respiratory movements, and stabilizes parts of the appendicular skeleton.
The Skull
The bones of the skull support the structures of the face and protect the brain. The skull consists of 22 bones, which are divided into two categories: cranial bones and facial bones. The cranial bones are eight bones that form the cranial cavity, which encloses the brain and serves as an attachment site for the muscles of the head and neck. The eight cranial bones are the frontal bone, two parietal bones, two temporal bones, occipital bone, sphenoid bone, and the ethmoid bone. Although the bones developed separately in the embryo and fetus, in the adult, they are tightly fused with connective tissue and adjoining bones do not move (Figure \(5\)).
The auditory ossicles of the middle ear transmit sounds from the air as vibrations to the fluid-filled cochlea. The auditory ossicles consist of six bones: two malleus bones, two incus bones, and two stapes on each side. These are the smallest bones in the body and are unique to mammals.
Fourteen facial bones form the face, provide cavities for the sense organs (eyes, mouth, and nose), protect the entrances to the digestive and respiratory tracts, and serve as attachment points for facial muscles. The 14 facial bones are the nasal bones, the maxillary bones, zygomatic bones, palatine, vomer, lacrimal bones, the inferior nasal conchae, and the mandible. All of these bones occur in pairs except for the mandible and the vomer (Figure \(6\)).
Although it is not found in the skull, the hyoid bone is considered a component of the axial skeleton. The hyoid bone lies below the mandible in the front of the neck. It acts as a movable base for the tongue and is connected to muscles of the jaw, larynx, and tongue. The mandible articulates with the base of the skull. The mandible controls the opening to the airway and gut. In animals with teeth, the mandible brings the surfaces of the teeth in contact with the maxillary teeth.
The Vertebral Column
The vertebral column, or spinal column, surrounds and protects the spinal cord, supports the head, and acts as an attachment point for the ribs and muscles of the back and neck. The adult vertebral column comprises 26 bones: the 24 vertebrae, the sacrum, and the coccyx bones. In the adult, the sacrum is typically composed of five vertebrae that fuse into one. The coccyx is typically 3–4 vertebrae that fuse into one. Around the age of 70, the sacrum and the coccyx may fuse together. We begin life with approximately 33 vertebrae, but as we grow, several vertebrae fuse together. The adult vertebrae are further divided into the 7 cervical vertebrae, 12 thoracic vertebrae, and 5 lumbar vertebrae (Figure \(7\)).
Each vertebral body has a large hole in the center through which the nerves of the spinal cord pass. There is also a notch on each side through which the spinal nerves, which serve the body at that level, can exit from the spinal cord. The vertebral column is approximately 71 cm (28 inches) in adult male humans and is curved, which can be seen from a side view. The names of the spinal curves correspond to the region of the spine in which they occur. The thoracic and sacral curves are concave (curve inwards relative to the front of the body) and the cervical and lumbar curves are convex (curve outwards relative to the front of the body). The arched curvature of the vertebral column increases its strength and flexibility, allowing it to absorb shocks like a spring (Figure \(7\)).
Intervertebral discs composed of fibrous cartilage lie between adjacent vertebral bodies from the second cervical vertebra to the sacrum. Each disc is part of a joint that allows for some movement of the spine and acts as a cushion to absorb shocks from movements such as walking and running. Intervertebral discs also act as ligaments to bind vertebrae together. The inner part of discs, the nucleus pulposus, hardens as people age and becomes less elastic. This loss of elasticity diminishes its ability to absorb shocks.
The Thoracic Cage
The thoracic cage, also known as the ribcage, is the skeleton of the chest, and consists of the ribs, sternum, thoracic vertebrae, and costal cartilages (Figure \(8\).8). The thoracic cage encloses and protects the organs of the thoracic cavity, including the heart and lungs. It also provides support for the shoulder girdles and upper limbs, and serves as the attachment point for the diaphragm, muscles of the back, chest, neck, and shoulders. Changes in the volume of the thorax enable breathing.
The sternum, or breastbone, is a long, flat bone located at the anterior of the chest. It is formed from three bones that fuse in the adult. The ribs are 12 pairs of long, curved bones that attach to the thoracic vertebrae and curve toward the front of the body, forming the ribcage. Costal cartilages connect the anterior ends of the ribs to the sternum, with the exception of rib pairs 11 and 12, which are free-floating ribs.
Human Appendicular Skeleton
The appendicular skeleton is composed of the bones of the upper limbs (which function to grasp and manipulate objects) and the lower limbs (which permit locomotion). It also includes the pectoral girdle, or shoulder girdle, that attaches the upper limbs to the body, and the pelvic girdle that attaches the lower limbs to the body (Figure \(9\)).
The Pectoral Girdle
The pectoral girdle bones provide the points of attachment of the upper limbs to the axial skeleton. The human pectoral girdle consists of the clavicle (or collarbone) in the anterior, and the scapula (or shoulder blades) in the posterior (Figure \(10\)).
The clavicles are S-shaped bones that position the arms on the body. The clavicles lie horizontally across the front of the thorax (chest) just above the first rib. These bones are fairly fragile and are susceptible to fractures. For example, a fall with the arms outstretched causes the force to be transmitted to the clavicles, which can break if the force is excessive. The clavicle articulates with the sternum and the scapula.
The scapulae are flat, triangular bones that are located at the back of the pectoral girdle. They support the muscles crossing the shoulder joint. A ridge, called the spine, runs across the back of the scapula and can easily be felt through the skin (Figure \(10\)). The spine of the scapula is a good example of a bony protrusion that facilitates a broad area of attachment for muscles to bone.
The Upper Limb
The upper limb contains 30 bones in three regions: the arm (shoulder to elbow), the forearm (ulna and radius), and the wrist and hand (Figure \(11\)).
An articulation is any place at which two bones are joined. The humerus is the largest and longest bone of the upper limb and the only bone of the arm. It articulates with the scapula at the shoulder and with the forearm at the elbow. The forearm extends from the elbow to the wrist and consists of two bones: the ulna and the radius. The radius is located along the lateral (thumb) side of the forearm and articulates with the humerus at the elbow. The ulna is located on the medial aspect (pinky-finger side) of the forearm. It is longer than the radius. The ulna articulates with the humerus at the elbow. The radius and ulna also articulate with the carpal bones and with each other, which in vertebrates enables a variable degree of rotation of the carpus with respect to the long axis of the limb. The hand includes the eight bones of the carpus (wrist), the five bones of the metacarpus (palm), and the 14 bones of the phalanges (digits). Each digit consists of three phalanges, except for the thumb, when present, which has only two.
The Pelvic Girdle
The pelvic girdle attaches to the lower limbs of the axial skeleton. Because it is responsible for bearing the weight of the body and for locomotion, the pelvic girdle is securely attached to the axial skeleton by strong ligaments. It also has deep sockets with robust ligaments to securely attach the femur to the body. The pelvic girdle is further strengthened by two large hip bones. In adults, the hip bones, or coxal bones, are formed by the fusion of three pairs of bones: the ilium, ischium, and pubis. The pelvis joins together in the anterior of the body at a joint called the pubic symphysis and with the bones of the sacrum at the posterior of the body.
The female pelvis is slightly different from the male pelvis. Over generations of evolution, females with a wider pubic angle and larger diameter pelvic canal reproduced more successfully. Therefore, their offspring also had pelvic anatomy that enabled successful childbirth (Figure \(12\)).
The Lower Limb
The lower limb consists of the thigh, the leg, and the foot. The bones of the lower limb are the femur (thigh bone), patella (kneecap), tibia and fibula (bones of the leg), tarsals (bones of the ankle), and metatarsals and phalanges (bones of the foot) (Figure \(13\)). The bones of the lower limbs are thicker and stronger than the bones of the upper limbs because of the need to support the entire weight of the body and the resulting forces from locomotion. In addition to evolutionary fitness, the bones of an individual will respond to forces exerted upon them.
The femur, or thighbone, is the longest, heaviest, and strongest bone in the body. The femur and pelvis form the hip joint at the proximal end. At the distal end, the femur, tibia, and patella form the knee joint. The patella, or kneecap, is a triangular bone that lies anterior to the knee joint. The patella is embedded in the tendon of the femoral extensors (quadriceps). It improves knee extension by reducing friction. The tibia, or shinbone, is a large bone of the leg that is located directly below the knee. The tibia articulates with the femur at its proximal end, with the fibula and the tarsal bones at its distal end. It is the second largest bone in the human body and is responsible for transmitting the weight of the body from the femur to the foot. The fibula, or calf bone, parallels and articulates with the tibia. It does not articulate with the femur and does not bear weight. The fibula acts as a site for muscle attachment and forms the lateral part of the ankle joint.
The tarsals are the seven bones of the ankle. The ankle transmits the weight of the body from the tibia and the fibula to the foot. The metatarsals are the five bones of the foot. The phalanges are the 14 bones of the toes. Each toe consists of three phalanges, except for the big toe that has only two (Figure \(14\)). Variations exist in other species; for example, the horse’s metacarpals and metatarsals are oriented vertically and do not make contact with the substrate.
Evolution Connection: Evolution of Body Design for Locomotion on Land
The transition of vertebrates onto land required a number of changes in body design, as movement on land presents a number of challenges for animals that are adapted to movement in water. The buoyancy of water provides a certain amount of lift, and a common form of movement by fish is lateral undulations of the entire body. This back and forth movement pushes the body against the water, creating forward movement. In most fish, the muscles of paired fins attach to girdles within the body, allowing for some control of locomotion. As certain fish began moving onto land, they retained their lateral undulation form of locomotion (anguilliform). However, instead of pushing against water, their fins or flippers became points of contact with the ground, around which they rotated their bodies.
The effect of gravity and the lack of buoyancy on land meant that body weight was suspended on the limbs, leading to increased strengthening and ossification of the limbs. The effect of gravity also required changes to the axial skeleton. Lateral undulations of land animal vertebral columns cause torsional strain. A firmer, more ossified vertebral column became common in terrestrial tetrapods because it reduces strain while providing the strength needed to support the body’s weight. In later tetrapods, the vertebrae began allowing for vertical motion rather than lateral flexion. Another change in the axial skeleton was the loss of a direct attachment between the pectoral girdle and the head. This reduced the jarring to the head caused by the impact of the limbs on the ground. The vertebrae of the neck also evolved to allow movement of the head independently of the body.
The appendicular skeleton of land animals is also different from aquatic animals. The shoulders attach to the pectoral girdle through muscles and connective tissue, thus reducing the jarring of the skull. Because of a lateral undulating vertebral column, in early tetrapods, the limbs were splayed out to the side and movement occurred by performing “push-ups.” The vertebrae of these animals had to move side-to-side in a similar manner to fish and reptiles. This type of motion requires large muscles to move the limbs toward the midline; it was almost like walking while doing push-ups, and it is not an efficient use of energy. Later tetrapods have their limbs placed under their bodies, so that each stride requires less force to move forward. This resulted in decreased adductor muscle size and an increased range of motion of the scapulae. This also restricts movement primarily to one plane, creating forward motion rather than moving the limbs upward as well as forward. The femur and humerus were also rotated, so that the ends of the limbs and digits were pointed forward, in the direction of motion, rather than out to the side. By placement underneath the body, limbs can swing forward like a pendulum to produce a stride that is more efficient for moving over land.
Summary
The three types of skeleton designs are hydrostatic skeletons, exoskeletons, and endoskeletons. A hydrostatic skeleton is formed by a fluid-filled compartment held under hydrostatic pressure; movement is created by the muscles producing pressure on the fluid. An exoskeleton is a hard external skeleton that protects the outer surface of an organism and enables movement through muscles attached on the inside. An endoskeleton is an internal skeleton composed of hard, mineralized tissue that also enables movement by attachment to muscles. The human skeleton is an endoskeleton that is composed of the axial and appendicular skeleton. The axial skeleton is composed of the bones of the skull, ossicles of the ear, hyoid bone, vertebral column, and ribcage. The skull consists of eight cranial bones and 14 facial bones. Six bones make up the ossicles of the middle ear, while the hyoid bone is located in the neck under the mandible. The vertebral column contains 26 bones, and it surrounds and protects the spinal cord. The thoracic cage consists of the sternum, ribs, thoracic vertebrae, and costal cartilages. The appendicular skeleton is made up of the limbs of the upper and lower limbs. The pectoral girdle is composed of the clavicles and the scapulae. The upper limb contains 30 bones in the arm, the forearm, and the hand. The pelvic girdle attaches the lower limbs to the axial skeleton. The lower limb includes the bones of the thigh, the leg, and the foot.
Glossary
appendicular skeleton
composed of the bones of the upper limbs, which function to grasp and manipulate objects, and the lower limbs, which permit locomotion
articulation
any place where two bones are joined
auditory ossicle
(also, middle ear) transduces sounds from the air into vibrations in the fluid-filled cochlea
axial skeleton
forms the central axis of the body and includes the bones of the skull, the ossicles of the middle ear, the hyoid bone of the throat, the vertebral column, and the thoracic cage (ribcage)
carpus
eight bones that comprise the wrist
clavicle
S-shaped bone that positions the arms laterally
coxal bone
hip bone
cranial bone
one of eight bones that form the cranial cavity that encloses the brain and serves as an attachment site for the muscles of the head and neck
endoskeleton
skeleton of living cells that produce a hard, mineralized tissue located within the soft tissue of organisms
exoskeleton
a secreted cellular product external skeleton that consists of a hard encasement on the surface of an organism
facial bone
one of the 14 bones that form the face; provides cavities for the sense organs (eyes, mouth, and nose) and attachment points for facial muscles
femur
(also, thighbone) longest, heaviest, and strongest bone in the body
fibula
(also, calf bone) parallels and articulates with the tibia
forearm
extends from the elbow to the wrist and consists of two bones: the ulna and the radius
humerus
only bone of the arm
hydrostatic skeleton
skeleton that consists of aqueous fluid held under pressure in a closed body compartment
hyoid bone
lies below the mandible in the front of the neck
intervertebral disc
composed of fibrous cartilage; lies between adjacent vertebrae from the second cervical vertebra to the sacrum
lower limb
consists of the thigh, the leg, and the foot
metacarpus
five bones that comprise the palm
metatarsal
one of the five bones of the foot
patella
(also, kneecap) triangular bone that lies anterior to the knee joint
pectoral girdle
bones that transmit the force generated by the upper limbs to the axial skeleton
phalange
one of the bones of the fingers or toes
pelvic girdle
bones that transmit the force generated by the lower limbs to the axial skeleton
radius
bone located along the lateral (thumb) side of the forearm; articulates with the humerus at the elbow
rib
one of 12 pairs of long, curved bones that attach to the thoracic vertebrae and curve toward the front of the body to form the ribcage
scapula
flat, triangular bone located at the posterior pectoral girdle
skull
bone that supports the structures of the face and protects the brain
sternum
(also, breastbone) long, flat bone located at the front of the chest
tarsal
one of the seven bones of the ankle
thoracic cage
(also, ribcage) skeleton of the chest, which consists of the ribs, thoracic vertebrae, sternum, and costal cartilages
tibia
(also, shinbone) large bone of the leg that is located directly below the knee
ulna
bone located on the medial aspect (pinky-finger side) of the forearm
vertebral column
(also, spine) surrounds and protects the spinal cord, supports the head, and acts as an attachment point for ribs and muscles of the back and neck | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/45%3A_The_Musculoskeletal_System/45.05%3A_Vertebrate_Skeleton_Evolution_and_Modes_of_Locomotion.txt |
• 46.1: Types of Digestive Systems
Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are processes that modify food within the animal body to make the nutrients and organic molecules needed for cellular function.
• 46.2: The Mouth and Teeth- Food Capture and Bulk Processing
Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are processes that modify food within the animal body to make the nutrients and organic molecules needed for cellular function.
• 46.3: The Esophagus and Stomach- The Early Stages of Digestion
Obtaining nutrition and energy from food is a multi-step process. For true animals, the first step is ingestion, the act of taking in food. This is followed by digestion, absorption, and elimination. In the following sections, each of these steps will be discussed in detail.
• 46.4: The Intestines- Breakdown, Absorption, and Elimination
Obtaining nutrition and energy from food is a multi-step process. For true animals, the first step is ingestion, the act of taking in food. This is followed by digestion, absorption, and elimination. In the following sections, each of these steps will be discussed in detail.
• 46.5: Accessory Organ Functions
• 46.6: Neural and Hormonal Regulation of the Digestive Tract
The brain is the control center for the sensation of hunger and satiety. The functions of the digestive system are regulated through neural and hormonal responses.
• 46.7: Food Energy, Energy Expenditure, and Essential Nutrients
• 46.8: Variations in Vertebrate Digestive Systems
Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are processes that modify food within the animal body to make the nutrients and organic molecules needed for cellular function.
46: The Digestive System
Skills to Develop
• Explain the processes of digestion and absorption
• Compare and contrast different types of digestive systems
• Explain the specialized functions of the organs involved in processing food in the body
• Describe the ways in which organs work together to digest food and absorb nutrients
Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are a number of processes that modify food within the animal body in order to make the nutrients and organic molecules accessible for cellular function. As animals evolved in complexity of form and function, their digestive systems have also evolved to accommodate their various dietary needs.
Herbivores, Omnivores, and Carnivores
Herbivores are animals whose primary food source is plant-based. Examples of herbivores, as shown in Figure \(1\) include vertebrates like deer, koalas, and some bird species, as well as invertebrates such as crickets and caterpillars. These animals have evolved digestive systems capable of handling large amounts of plant material. Herbivores can be further classified into frugivores (fruit-eaters), granivores (seed eaters), nectivores (nectar feeders), and folivores (leaf eaters).
Carnivores are animals that eat other animals. The word carnivore is derived from Latin and literally means “meat eater.” Wild cats such as lions, shown in Figure \(2\)a and tigers are examples of vertebrate carnivores, as are snakes and sharks, while invertebrate carnivores include sea stars, spiders, and ladybugs, shown in Figure \(2\)b. Obligate carnivores are those that rely entirely on animal flesh to obtain their nutrients; examples of obligate carnivores are members of the cat family, such as lions and cheetahs. Facultative carnivores are those that also eat non-animal food in addition to animal food. Note that there is no clear line that differentiates facultative carnivores from omnivores; dogs would be considered facultative carnivores.
Omnivores are animals that eat both plant- and animal-derived food. In Latin, omnivore means to eat everything. Humans, bears (shown in Figure \(3\)a), and chickens are example of vertebrate omnivores; invertebrate omnivores include cockroaches and crayfish (shown in Figure \(3\)b).
Invertebrate Digestive Systems
Animals have evolved different types of digestive systems to aid in the digestion of the different foods they consume. The simplest example is that of a gastrovascular cavity and is found in organisms with only one opening for digestion. Platyhelminthes (flatworms), Ctenophora (comb jellies), and Cnidaria (coral, jelly fish, and sea anemones) use this type of digestion. Gastrovascular cavities, as shown in Figure \(4\)a, are typically a blind tube or cavity with only one opening, the “mouth”, which also serves as an “anus”. Ingested material enters the mouth and passes through a hollow, tubular cavity. Cells within the cavity secrete digestive enzymes that break down the food. The food particles are engulfed by the cells lining the gastrovascular cavity.
The alimentary canal, shown in Figure \(4\)b, is a more advanced system: it consists of one tube with a mouth at one end and an anus at the other. Earthworms are an example of an animal with an alimentary canal. Once the food is ingested through the mouth, it passes through the esophagus and is stored in an organ called the crop; then it passes into the gizzard where it is churned and digested. From the gizzard, the food passes through the intestine, the nutrients are absorbed, and the waste is eliminated as feces, called castings, through the anus.
Vertebrate Digestive Systems
Vertebrates have evolved more complex digestive systems to adapt to their dietary needs. Some animals have a single stomach, while others have multi-chambered stomachs. Birds have developed a digestive system adapted to eating unmasticated food.
Monogastric: Single-chambered Stomach
As the word monogastric suggests, this type of digestive system consists of one (“mono”) stomach chamber (“gastric”). Humans and many animals have a monogastric digestive system as illustrated in Figure \(5\). The process of digestion begins with the mouth and the intake of food. The teeth play an important role in masticating (chewing) or physically breaking down food into smaller particles. The enzymes present in saliva also begin to chemically break down food. The esophagus is a long tube that connects the mouth to the stomach. Using peristalsis, or wave-like smooth muscle contractions, the muscles of the esophagus push the food towards the stomach. In order to speed up the actions of enzymes in the stomach, the stomach is an extremely acidic environment, with a pH between 1.5 and 2.5. The gastric juices, which include enzymes in the stomach, act on the food particles and continue the process of digestion. Further breakdown of food takes place in the small intestine where enzymes produced by the liver, the small intestine, and the pancreas continue the process of digestion. The nutrients are absorbed into the blood stream across the epithelial cells lining the walls of the small intestines. The waste material travels on to the large intestine where water is absorbed and the drier waste material is compacted into feces; it is stored until it is excreted through the rectum.
Avian
Birds face special challenges when it comes to obtaining nutrition from food. They do not have teeth and so their digestive system, shown in Figure \(6\), must be able to process un-masticated food. Birds have evolved a variety of beak types that reflect the vast variety in their diet, ranging from seeds and insects to fruits and nuts. Because most birds fly, their metabolic rates are high in order to efficiently process food and keep their body weight low. The stomach of birds has two chambers: the proventriculus, where gastric juices are produced to digest the food before it enters the stomach, and the gizzard, where the food is stored, soaked, and mechanically ground. The undigested material forms food pellets that are sometimes regurgitated. Most of the chemical digestion and absorption happens in the intestine and the waste is excreted through the cloaca.
Evolution Connection: Avian Adaptations
Birds have a highly efficient, simplified digestive system. Recent fossil evidence has shown that the evolutionary divergence of birds from other land animals was characterized by streamlining and simplifying the digestive system. Unlike many other animals, birds do not have teeth to chew their food. In place of lips, they have sharp pointy beaks. The horny beak, lack of jaws, and the smaller tongue of the birds can be traced back to their dinosaur ancestors. The emergence of these changes seems to coincide with the inclusion of seeds in the bird diet. Seed-eating birds have beaks that are shaped for grabbing seeds and the two-compartment stomach allows for delegation of tasks. Since birds need to remain light in order to fly, their metabolic rates are very high, which means they digest their food very quickly and need to eat often. Contrast this with the ruminants, where the digestion of plant matter takes a very long time.
Ruminants
Ruminants are mainly herbivores like cows, sheep, and goats, whose entire diet consists of eating large amounts of roughage or fiber. They have evolved digestive systems that help them digest vast amounts of cellulose. An interesting feature of the ruminants’ mouth is that they do not have upper incisor teeth. They use their lower teeth, tongue and lips to tear and chew their food. From the mouth, the food travels to the esophagus and on to the stomach.
To help digest the large amount of plant material, the stomach of the ruminants is a multi-chambered organ, as illustrated in Figure \(7\). The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that break down cellulose and ferment ingested food. The abomasum is the “true” stomach and is the equivalent of the monogastric stomach chamber where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the microbial support necessary to digest plant material in ruminants. The fermentation process produces large amounts of gas in the stomach chamber, which must be eliminated. As in other animals, the small intestine plays an important role in nutrient absorption, and the large intestine helps in the elimination of waste.
Pseudo-ruminants
Some animals, such as camels and alpacas, are pseudo-ruminants. They eat a lot of plant material and roughage. Digesting plant material is not easy because plant cell walls contain the polymeric sugar molecule cellulose. The digestive enzymes of these animals cannot break down cellulose, but microorganisms present in the digestive system can. Therefore, the digestive system must be able to handle large amounts of roughage and break down the cellulose. Pseudo-ruminants have a three-chamber stomach in the digestive system. However, their cecum—a pouched organ at the beginning of the large intestine containing many microorganisms that are necessary for the digestion of plant materials—is large and is the site where the roughage is fermented and digested. These animals do not have a rumen but have an omasum, abomasum, and reticulum.
Parts of the Digestive System
The vertebrate digestive system is designed to facilitate the transformation of food matter into the nutrient components that sustain organisms.
Oral Cavity
The oral cavity, or mouth, is the point of entry of food into the digestive system, illustrated in Figure \(8\). The food consumed is broken into smaller particles by mastication, the chewing action of the teeth. All mammals have teeth and can chew their food.
The extensive chemical process of digestion begins in the mouth. As food is being chewed, saliva, produced by the salivary glands, mixes with the food. Saliva is a watery substance produced in the mouths of many animals. There are three major glands that secrete saliva—the parotid, the submandibular, and the sublingual. Saliva contains mucus that moistens food and buffers the pH of the food. Saliva also contains immunoglobulins and lysozymes, which have antibacterial action to reduce tooth decay by inhibiting growth of some bacteria. Saliva also contains an enzyme called salivary amylase that begins the process of converting starches in the food into a disaccharide called maltose. Another enzyme called lipase is produced by the cells in the tongue. Lipases are a class of enzymes that can break down triglycerides. The lingual lipase begins the breakdown of fat components in the food. The chewing and wetting action provided by the teeth and saliva prepare the food into a mass called the bolus for swallowing. The tongue helps in swallowing—moving the bolus from the mouth into the pharynx. The pharynx opens to two passageways: the trachea, which leads to the lungs, and the esophagus, which leads to the stomach. The trachea has an opening called the glottis, which is covered by a cartilaginous flap called the epiglottis. When swallowing, the epiglottis closes the glottis and food passes into the esophagus and not the trachea. This arrangement allows food to be kept out of the trachea.
Esophagus
The esophagus is a tubular organ that connects the mouth to the stomach. The chewed and softened food passes through the esophagus after being swallowed. The smooth muscles of the esophagus undergo a series of wave like movements called peristalsis that push the food toward the stomach, as illustrated in Figure \(9\). The peristalsis wave is unidirectional—it moves food from the mouth to the stomach, and reverse movement is not possible. The peristaltic movement of the esophagus is an involuntary reflex; it takes place in response to the act of swallowing.
A ring-like muscle called a sphincter forms valves in the digestive system. The gastro-esophageal sphincter is located at the stomach end of the esophagus. In response to swallowing and the pressure exerted by the bolus of food, this sphincter opens, and the bolus enters the stomach. When there is no swallowing action, this sphincter is shut and prevents the contents of the stomach from traveling up the esophagus. Many animals have a true sphincter; however, in humans, there is no true sphincter, but the esophagus remains closed when there is no swallowing action. Acid reflux or “heartburn” occurs when the acidic digestive juices escape into the esophagus.
Stomach
A large part of digestion occurs in the stomach, shown in Figure \(10\). The stomach is a saclike organ that secretes gastric digestive juices. The pH in the stomach is between 1.5 and 2.5. This highly acidic environment is required for the chemical breakdown of food and the extraction of nutrients. When empty, the stomach is a rather small organ; however, it can expand to up to 20 times its resting size when filled with food. This characteristic is particularly useful for animals that need to eat when food is available.
Art Connection
Which of the following statements about the digestive system is false?
1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.
The stomach is also the major site for protein digestion in animals other than ruminants. Protein digestion is mediated by an enzyme called pepsin in the stomach chamber. Pepsin is secreted by the chief cells in the stomach in an inactive form called pepsinogen. Pepsin breaks peptide bonds and cleaves proteins into smaller polypeptides; it also helps activate more pepsinogen, starting a positive feedback mechanism that generates more pepsin. Another cell type—parietal cells—secrete hydrogen and chloride ions, which combine in the lumen to form hydrochloric acid, the primary acidic component of the stomach juices. Hydrochloric acid helps to convert the inactive pepsinogen to pepsin. The highly acidic environment also kills many microorganisms in the food and, combined with the action of the enzyme pepsin, results in the hydrolysis of protein in the food. Chemical digestion is facilitated by the churning action of the stomach. Contraction and relaxation of smooth muscles mixes the stomach contents about every 20 minutes. The partially digested food and gastric juice mixture is called chyme. Chyme passes from the stomach to the small intestine. Further protein digestion takes place in the small intestine. Gastric emptying occurs within two to six hours after a meal. Only a small amount of chyme is released into the small intestine at a time. The movement of chyme from the stomach into the small intestine is regulated by the pyloric sphincter.
When digesting protein and some fats, the stomach lining must be protected from getting digested by pepsin. There are two points to consider when describing how the stomach lining is protected. First, as previously mentioned, the enzyme pepsin is synthesized in the inactive form. This protects the chief cells, because pepsinogen does not have the same enzyme functionality of pepsin. Second, the stomach has a thick mucus lining that protects the underlying tissue from the action of the digestive juices. When this mucus lining is ruptured, ulcers can form in the stomach. Ulcers are open wounds in or on an organ caused by bacteria (Helicobacter pylori) when the mucus lining is ruptured and fails to reform.
Small Intestine
Chyme moves from the stomach to the small intestine. The small intestine is the organ where the digestion of protein, fats, and carbohydrates is completed. The small intestine is a long tube-like organ with a highly folded surface containing finger-like projections called the villi. The apical surface of each villus has many microscopic projections called microvilli. These structures, illustrated in Figure \(11\), are lined with epithelial cells on the luminal side and allow for the nutrients to be absorbed from the digested food and absorbed into the blood stream on the other side. The villi and microvilli, with their many folds, increase the surface area of the intestine and increase absorption efficiency of the nutrients. Absorbed nutrients in the blood are carried into the hepatic portal vein, which leads to the liver. There, the liver regulates the distribution of nutrients to the rest of the body and removes toxic substances, including drugs, alcohol, and some pathogens.
Art Connection
Which of the following statements about the small intestine is false?
1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.
The human small intestine is over 6m long and is divided into three parts: the duodenum, the jejunum, and the ileum. The “C-shaped,” fixed part of the small intestine is called the duodenum and is shown in Figure \(11\). The duodenum is separated from the stomach by the pyloric sphincter which opens to allow chyme to move from the stomach to the duodenum. In the duodenum, chyme is mixed with pancreatic juices in an alkaline solution rich in bicarbonate that neutralizes the acidity of chyme and acts as a buffer. Pancreatic juices also contain several digestive enzymes. Digestive juices from the pancreas, liver, and gallbladder, as well as from gland cells of the intestinal wall itself, enter the duodenum. Bile is produced in the liver and stored and concentrated in the gallbladder. Bile contains bile salts which emulsify lipids while the pancreas produces enzymes that catabolize starches, disaccharides, proteins, and fats. These digestive juices break down the food particles in the chyme into glucose, triglycerides, and amino acids. Some chemical digestion of food takes place in the duodenum. Absorption of fatty acids also takes place in the duodenum.
The second part of the small intestine is called the jejunum, shown in Figure \(11\). Here, hydrolysis of nutrients is continued while most of the carbohydrates and amino acids are absorbed through the intestinal lining. The bulk of chemical digestion and nutrient absorption occurs in the jejunum.
The ileum, also illustrated in Figure \(11\) is the last part of the small intestine and here the bile salts and vitamins are absorbed into blood stream. The undigested food is sent to the colon from the ileum via peristaltic movements of the muscle. The ileum ends and the large intestine begins at the ileocecal valve. The vermiform, “worm-like,” appendix is located at the ileocecal valve. The appendix of humans secretes no enzymes and has an insignificant role in immunity.
Large Intestine
The large intestine, illustrated in Figure \(12\), reabsorbs the water from the undigested food material and processes the waste material. The human large intestine is much smaller in length compared to the small intestine but larger in diameter. It has three parts: the cecum, the colon, and the rectum. The cecum joins the ileum to the colon and is the receiving pouch for the waste matter. The colon is home to many bacteria or “intestinal flora” that aid in the digestive processes. The colon can be divided into four regions, the ascending colon, the transverse colon, the descending colon and the sigmoid colon. The main functions of the colon are to extract the water and mineral salts from undigested food, and to store waste material. Carnivorous mammals have a shorter large intestine compared to herbivorous mammals due to their diet.
Rectum and Anus
The rectum is the terminal end of the large intestine, as shown in Figure \(12\). The primary role of the rectum is to store the feces until defecation. The feces are propelled using peristaltic movements during elimination. The anus is an opening at the far-end of the digestive tract and is the exit point for the waste material. Two sphincters between the rectum and anus control elimination: the inner sphincter is involuntary and the outer sphincter is voluntary.
Accessory Organs
The organs discussed above are the organs of the digestive tract through which food passes. Accessory organs are organs that add secretions (enzymes) that catabolize food into nutrients. Accessory organs include salivary glands, the liver, the pancreas, and the gallbladder. The liver, pancreas, and gallbladder are regulated by hormones in response to the food consumed.
The liver is the largest internal organ in humans and it plays a very important role in digestion of fats and detoxifying blood. The liver produces bile, a digestive juice that is required for the breakdown of fatty components of the food in the duodenum. The liver also processes the vitamins and fats and synthesizes many plasma proteins.
The pancreas is another important gland that secretes digestive juices. The chyme produced from the stomach is highly acidic in nature; the pancreatic juices contain high levels of bicarbonate, an alkali that neutralizes the acidic chyme. Additionally, the pancreatic juices contain a large variety of enzymes that are required for the digestion of protein and carbohydrates.
The gallbladder is a small organ that aids the liver by storing bile and concentrating bile salts. When chyme containing fatty acids enters the duodenum, the bile is secreted from the gallbladder into the duodenum.
Summary
Different animals have evolved different types of digestive systems specialized to meet their dietary needs. Humans and many other animals have monogastric digestive systems with a single-chambered stomach. Birds have evolved a digestive system that includes a gizzard where the food is crushed into smaller pieces. This compensates for their inability to masticate. Ruminants that consume large amounts of plant material have a multi-chambered stomach that digests roughage. Pseudo-ruminants have similar digestive processes as ruminants but do not have the four-compartment stomach. Processing food involves ingestion (eating), digestion (mechanical and enzymatic breakdown of large molecules), absorption (cellular uptake of nutrients), and elimination (removal of undigested waste as feces).
Many organs work together to digest food and absorb nutrients. The mouth is the point of ingestion and the location where both mechanical and chemical breakdown of food begins. Saliva contains an enzyme called amylase that breaks down carbohydrates. The food bolus travels through the esophagus by peristaltic movements to the stomach. The stomach has an extremely acidic environment. An enzyme called pepsin digests protein in the stomach. Further digestion and absorption take place in the small intestine. The large intestine reabsorbs water from the undigested food and stores waste until elimination.
Art Connections
Figure \(10\): Which of the following statements about the digestive system is false?
1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.
Answer
B
Figure \(11\): Which of the following statements about the small intestine is false?
1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.
Answer
C
Glossary
alimentary canal
tubular digestive system with a mouth and anus
anus
exit point for waste material
bile
digestive juice produced by the liver; important for digestion of lipids
bolus
mass of food resulting from chewing action and wetting by saliva
carnivore
animal that consumes animal flesh
chyme
mixture of partially digested food and stomach juices
duodenum
first part of the small intestine where a large part of digestion of carbohydrates and fats occurs
esophagus
tubular organ that connects the mouth to the stomach
gallbladder
organ that stores and concentrates bile
gastrovascular cavity
digestive system consisting of a single opening
gizzard
muscular organ that grinds food
herbivore
animal that consumes strictly plant diet
ileum
last part of the small intestine; connects the small intestine to the large intestine; important for absorption of B-12
jejunum
second part of the small intestine
large intestine
digestive system organ that reabsorbs water from undigested material and processes waste matter
lipase
enzyme that chemically breaks down lipids
liver
organ that produces bile for digestion and processes vitamins and lipids
monogastric
digestive system that consists of a single-chambered stomach
omnivore
animal that consumes both plants and animals
pancreas
gland that secretes digestive juices
pepsin
enzyme found in the stomach whose main role is protein digestion
pepsinogen
inactive form of pepsin
peristalsis
wave-like movements of muscle tissue
proventriculus
glandular part of a bird’s stomach
rectum
area of the body where feces is stored until elimination
roughage
component of food that is low in energy and high in fiber
ruminant
animal with a stomach divided into four compartments
salivary amylase
enzyme found in saliva, which converts carbohydrates to maltose
small intestine
organ where digestion of protein, fats, and carbohydrates is completed
sphincter
band of muscle that controls movement of materials throughout the digestive tract
stomach
saclike organ containing acidic digestive juices
villi
folds on the inner surface of the small intestine whose role is to increase absorption area | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/46%3A_The_Digestive_System/46.01%3A_Types_of_Digestive_Systems.txt |
Skills to Develop
• Explain the processes of digestion and absorption
• Compare and contrast different types of digestive systems
• Explain the specialized functions of the organs involved in processing food in the body
• Describe the ways in which organs work together to digest food and absorb nutrients
Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are a number of processes that modify food within the animal body in order to make the nutrients and organic molecules accessible for cellular function. As animals evolved in complexity of form and function, their digestive systems have also evolved to accommodate their various dietary needs.
Herbivores, Omnivores, and Carnivores
Herbivores are animals whose primary food source is plant-based. Examples of herbivores, as shown in Figure \(1\) include vertebrates like deer, koalas, and some bird species, as well as invertebrates such as crickets and caterpillars. These animals have evolved digestive systems capable of handling large amounts of plant material. Herbivores can be further classified into frugivores (fruit-eaters), granivores (seed eaters), nectivores (nectar feeders), and folivores (leaf eaters).
Carnivores are animals that eat other animals. The word carnivore is derived from Latin and literally means “meat eater.” Wild cats such as lions, shown in Figure \(2\)a and tigers are examples of vertebrate carnivores, as are snakes and sharks, while invertebrate carnivores include sea stars, spiders, and ladybugs, shown in Figure \(2\)b. Obligate carnivores are those that rely entirely on animal flesh to obtain their nutrients; examples of obligate carnivores are members of the cat family, such as lions and cheetahs. Facultative carnivores are those that also eat non-animal food in addition to animal food. Note that there is no clear line that differentiates facultative carnivores from omnivores; dogs would be considered facultative carnivores.
Omnivores are animals that eat both plant- and animal-derived food. In Latin, omnivore means to eat everything. Humans, bears (shown in Figure \(3\)a), and chickens are example of vertebrate omnivores; invertebrate omnivores include cockroaches and crayfish (shown in Figure \(3\)b).
Invertebrate Digestive Systems
Animals have evolved different types of digestive systems to aid in the digestion of the different foods they consume. The simplest example is that of a gastrovascular cavity and is found in organisms with only one opening for digestion. Platyhelminthes (flatworms), Ctenophora (comb jellies), and Cnidaria (coral, jelly fish, and sea anemones) use this type of digestion. Gastrovascular cavities, as shown in Figure \(4\)a, are typically a blind tube or cavity with only one opening, the “mouth”, which also serves as an “anus”. Ingested material enters the mouth and passes through a hollow, tubular cavity. Cells within the cavity secrete digestive enzymes that break down the food. The food particles are engulfed by the cells lining the gastrovascular cavity.
The alimentary canal, shown in Figure \(4\)b, is a more advanced system: it consists of one tube with a mouth at one end and an anus at the other. Earthworms are an example of an animal with an alimentary canal. Once the food is ingested through the mouth, it passes through the esophagus and is stored in an organ called the crop; then it passes into the gizzard where it is churned and digested. From the gizzard, the food passes through the intestine, the nutrients are absorbed, and the waste is eliminated as feces, called castings, through the anus.
Vertebrate Digestive Systems
Vertebrates have evolved more complex digestive systems to adapt to their dietary needs. Some animals have a single stomach, while others have multi-chambered stomachs. Birds have developed a digestive system adapted to eating unmasticated food.
Monogastric: Single-chambered Stomach
As the word monogastric suggests, this type of digestive system consists of one (“mono”) stomach chamber (“gastric”). Humans and many animals have a monogastric digestive system as illustrated in Figure \(5\). The process of digestion begins with the mouth and the intake of food. The teeth play an important role in masticating (chewing) or physically breaking down food into smaller particles. The enzymes present in saliva also begin to chemically break down food. The esophagus is a long tube that connects the mouth to the stomach. Using peristalsis, or wave-like smooth muscle contractions, the muscles of the esophagus push the food towards the stomach. In order to speed up the actions of enzymes in the stomach, the stomach is an extremely acidic environment, with a pH between 1.5 and 2.5. The gastric juices, which include enzymes in the stomach, act on the food particles and continue the process of digestion. Further breakdown of food takes place in the small intestine where enzymes produced by the liver, the small intestine, and the pancreas continue the process of digestion. The nutrients are absorbed into the blood stream across the epithelial cells lining the walls of the small intestines. The waste material travels on to the large intestine where water is absorbed and the drier waste material is compacted into feces; it is stored until it is excreted through the rectum.
Avian
Birds face special challenges when it comes to obtaining nutrition from food. They do not have teeth and so their digestive system, shown in Figure \(6\), must be able to process un-masticated food. Birds have evolved a variety of beak types that reflect the vast variety in their diet, ranging from seeds and insects to fruits and nuts. Because most birds fly, their metabolic rates are high in order to efficiently process food and keep their body weight low. The stomach of birds has two chambers: the proventriculus, where gastric juices are produced to digest the food before it enters the stomach, and the gizzard, where the food is stored, soaked, and mechanically ground. The undigested material forms food pellets that are sometimes regurgitated. Most of the chemical digestion and absorption happens in the intestine and the waste is excreted through the cloaca.
Evolution Connection: Avian Adaptations
Birds have a highly efficient, simplified digestive system. Recent fossil evidence has shown that the evolutionary divergence of birds from other land animals was characterized by streamlining and simplifying the digestive system. Unlike many other animals, birds do not have teeth to chew their food. In place of lips, they have sharp pointy beaks. The horny beak, lack of jaws, and the smaller tongue of the birds can be traced back to their dinosaur ancestors. The emergence of these changes seems to coincide with the inclusion of seeds in the bird diet. Seed-eating birds have beaks that are shaped for grabbing seeds and the two-compartment stomach allows for delegation of tasks. Since birds need to remain light in order to fly, their metabolic rates are very high, which means they digest their food very quickly and need to eat often. Contrast this with the ruminants, where the digestion of plant matter takes a very long time.
Ruminants
Ruminants are mainly herbivores like cows, sheep, and goats, whose entire diet consists of eating large amounts of roughage or fiber. They have evolved digestive systems that help them digest vast amounts of cellulose. An interesting feature of the ruminants’ mouth is that they do not have upper incisor teeth. They use their lower teeth, tongue and lips to tear and chew their food. From the mouth, the food travels to the esophagus and on to the stomach.
To help digest the large amount of plant material, the stomach of the ruminants is a multi-chambered organ, as illustrated in Figure \(7\). The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that break down cellulose and ferment ingested food. The abomasum is the “true” stomach and is the equivalent of the monogastric stomach chamber where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the microbial support necessary to digest plant material in ruminants. The fermentation process produces large amounts of gas in the stomach chamber, which must be eliminated. As in other animals, the small intestine plays an important role in nutrient absorption, and the large intestine helps in the elimination of waste.
Pseudo-ruminants
Some animals, such as camels and alpacas, are pseudo-ruminants. They eat a lot of plant material and roughage. Digesting plant material is not easy because plant cell walls contain the polymeric sugar molecule cellulose. The digestive enzymes of these animals cannot break down cellulose, but microorganisms present in the digestive system can. Therefore, the digestive system must be able to handle large amounts of roughage and break down the cellulose. Pseudo-ruminants have a three-chamber stomach in the digestive system. However, their cecum—a pouched organ at the beginning of the large intestine containing many microorganisms that are necessary for the digestion of plant materials—is large and is the site where the roughage is fermented and digested. These animals do not have a rumen but have an omasum, abomasum, and reticulum.
Parts of the Digestive System
The vertebrate digestive system is designed to facilitate the transformation of food matter into the nutrient components that sustain organisms.
Oral Cavity
The oral cavity, or mouth, is the point of entry of food into the digestive system, illustrated in Figure \(8\). The food consumed is broken into smaller particles by mastication, the chewing action of the teeth. All mammals have teeth and can chew their food.
The extensive chemical process of digestion begins in the mouth. As food is being chewed, saliva, produced by the salivary glands, mixes with the food. Saliva is a watery substance produced in the mouths of many animals. There are three major glands that secrete saliva—the parotid, the submandibular, and the sublingual. Saliva contains mucus that moistens food and buffers the pH of the food. Saliva also contains immunoglobulins and lysozymes, which have antibacterial action to reduce tooth decay by inhibiting growth of some bacteria. Saliva also contains an enzyme called salivary amylase that begins the process of converting starches in the food into a disaccharide called maltose. Another enzyme called lipase is produced by the cells in the tongue. Lipases are a class of enzymes that can break down triglycerides. The lingual lipase begins the breakdown of fat components in the food. The chewing and wetting action provided by the teeth and saliva prepare the food into a mass called the bolus for swallowing. The tongue helps in swallowing—moving the bolus from the mouth into the pharynx. The pharynx opens to two passageways: the trachea, which leads to the lungs, and the esophagus, which leads to the stomach. The trachea has an opening called the glottis, which is covered by a cartilaginous flap called the epiglottis. When swallowing, the epiglottis closes the glottis and food passes into the esophagus and not the trachea. This arrangement allows food to be kept out of the trachea.
Esophagus
The esophagus is a tubular organ that connects the mouth to the stomach. The chewed and softened food passes through the esophagus after being swallowed. The smooth muscles of the esophagus undergo a series of wave like movements called peristalsis that push the food toward the stomach, as illustrated in Figure \(9\). The peristalsis wave is unidirectional—it moves food from the mouth to the stomach, and reverse movement is not possible. The peristaltic movement of the esophagus is an involuntary reflex; it takes place in response to the act of swallowing.
A ring-like muscle called a sphincter forms valves in the digestive system. The gastro-esophageal sphincter is located at the stomach end of the esophagus. In response to swallowing and the pressure exerted by the bolus of food, this sphincter opens, and the bolus enters the stomach. When there is no swallowing action, this sphincter is shut and prevents the contents of the stomach from traveling up the esophagus. Many animals have a true sphincter; however, in humans, there is no true sphincter, but the esophagus remains closed when there is no swallowing action. Acid reflux or “heartburn” occurs when the acidic digestive juices escape into the esophagus.
Stomach
A large part of digestion occurs in the stomach, shown in Figure \(10\). The stomach is a saclike organ that secretes gastric digestive juices. The pH in the stomach is between 1.5 and 2.5. This highly acidic environment is required for the chemical breakdown of food and the extraction of nutrients. When empty, the stomach is a rather small organ; however, it can expand to up to 20 times its resting size when filled with food. This characteristic is particularly useful for animals that need to eat when food is available.
Art Connection
Which of the following statements about the digestive system is false?
1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.
The stomach is also the major site for protein digestion in animals other than ruminants. Protein digestion is mediated by an enzyme called pepsin in the stomach chamber. Pepsin is secreted by the chief cells in the stomach in an inactive form called pepsinogen. Pepsin breaks peptide bonds and cleaves proteins into smaller polypeptides; it also helps activate more pepsinogen, starting a positive feedback mechanism that generates more pepsin. Another cell type—parietal cells—secrete hydrogen and chloride ions, which combine in the lumen to form hydrochloric acid, the primary acidic component of the stomach juices. Hydrochloric acid helps to convert the inactive pepsinogen to pepsin. The highly acidic environment also kills many microorganisms in the food and, combined with the action of the enzyme pepsin, results in the hydrolysis of protein in the food. Chemical digestion is facilitated by the churning action of the stomach. Contraction and relaxation of smooth muscles mixes the stomach contents about every 20 minutes. The partially digested food and gastric juice mixture is called chyme. Chyme passes from the stomach to the small intestine. Further protein digestion takes place in the small intestine. Gastric emptying occurs within two to six hours after a meal. Only a small amount of chyme is released into the small intestine at a time. The movement of chyme from the stomach into the small intestine is regulated by the pyloric sphincter.
When digesting protein and some fats, the stomach lining must be protected from getting digested by pepsin. There are two points to consider when describing how the stomach lining is protected. First, as previously mentioned, the enzyme pepsin is synthesized in the inactive form. This protects the chief cells, because pepsinogen does not have the same enzyme functionality of pepsin. Second, the stomach has a thick mucus lining that protects the underlying tissue from the action of the digestive juices. When this mucus lining is ruptured, ulcers can form in the stomach. Ulcers are open wounds in or on an organ caused by bacteria (Helicobacter pylori) when the mucus lining is ruptured and fails to reform.
Small Intestine
Chyme moves from the stomach to the small intestine. The small intestine is the organ where the digestion of protein, fats, and carbohydrates is completed. The small intestine is a long tube-like organ with a highly folded surface containing finger-like projections called the villi. The apical surface of each villus has many microscopic projections called microvilli. These structures, illustrated in Figure \(11\), are lined with epithelial cells on the luminal side and allow for the nutrients to be absorbed from the digested food and absorbed into the blood stream on the other side. The villi and microvilli, with their many folds, increase the surface area of the intestine and increase absorption efficiency of the nutrients. Absorbed nutrients in the blood are carried into the hepatic portal vein, which leads to the liver. There, the liver regulates the distribution of nutrients to the rest of the body and removes toxic substances, including drugs, alcohol, and some pathogens.
Art Connection
Which of the following statements about the small intestine is false?
1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.
The human small intestine is over 6m long and is divided into three parts: the duodenum, the jejunum, and the ileum. The “C-shaped,” fixed part of the small intestine is called the duodenum and is shown in Figure \(11\). The duodenum is separated from the stomach by the pyloric sphincter which opens to allow chyme to move from the stomach to the duodenum. In the duodenum, chyme is mixed with pancreatic juices in an alkaline solution rich in bicarbonate that neutralizes the acidity of chyme and acts as a buffer. Pancreatic juices also contain several digestive enzymes. Digestive juices from the pancreas, liver, and gallbladder, as well as from gland cells of the intestinal wall itself, enter the duodenum. Bile is produced in the liver and stored and concentrated in the gallbladder. Bile contains bile salts which emulsify lipids while the pancreas produces enzymes that catabolize starches, disaccharides, proteins, and fats. These digestive juices break down the food particles in the chyme into glucose, triglycerides, and amino acids. Some chemical digestion of food takes place in the duodenum. Absorption of fatty acids also takes place in the duodenum.
The second part of the small intestine is called the jejunum, shown in Figure \(11\). Here, hydrolysis of nutrients is continued while most of the carbohydrates and amino acids are absorbed through the intestinal lining. The bulk of chemical digestion and nutrient absorption occurs in the jejunum.
The ileum, also illustrated in Figure \(11\) is the last part of the small intestine and here the bile salts and vitamins are absorbed into blood stream. The undigested food is sent to the colon from the ileum via peristaltic movements of the muscle. The ileum ends and the large intestine begins at the ileocecal valve. The vermiform, “worm-like,” appendix is located at the ileocecal valve. The appendix of humans secretes no enzymes and has an insignificant role in immunity.
Large Intestine
The large intestine, illustrated in Figure \(12\), reabsorbs the water from the undigested food material and processes the waste material. The human large intestine is much smaller in length compared to the small intestine but larger in diameter. It has three parts: the cecum, the colon, and the rectum. The cecum joins the ileum to the colon and is the receiving pouch for the waste matter. The colon is home to many bacteria or “intestinal flora” that aid in the digestive processes. The colon can be divided into four regions, the ascending colon, the transverse colon, the descending colon and the sigmoid colon. The main functions of the colon are to extract the water and mineral salts from undigested food, and to store waste material. Carnivorous mammals have a shorter large intestine compared to herbivorous mammals due to their diet.
Rectum and Anus
The rectum is the terminal end of the large intestine, as shown in Figure \(12\). The primary role of the rectum is to store the feces until defecation. The feces are propelled using peristaltic movements during elimination. The anus is an opening at the far-end of the digestive tract and is the exit point for the waste material. Two sphincters between the rectum and anus control elimination: the inner sphincter is involuntary and the outer sphincter is voluntary.
Accessory Organs
The organs discussed above are the organs of the digestive tract through which food passes. Accessory organs are organs that add secretions (enzymes) that catabolize food into nutrients. Accessory organs include salivary glands, the liver, the pancreas, and the gallbladder. The liver, pancreas, and gallbladder are regulated by hormones in response to the food consumed.
The liver is the largest internal organ in humans and it plays a very important role in digestion of fats and detoxifying blood. The liver produces bile, a digestive juice that is required for the breakdown of fatty components of the food in the duodenum. The liver also processes the vitamins and fats and synthesizes many plasma proteins.
The pancreas is another important gland that secretes digestive juices. The chyme produced from the stomach is highly acidic in nature; the pancreatic juices contain high levels of bicarbonate, an alkali that neutralizes the acidic chyme. Additionally, the pancreatic juices contain a large variety of enzymes that are required for the digestion of protein and carbohydrates.
The gallbladder is a small organ that aids the liver by storing bile and concentrating bile salts. When chyme containing fatty acids enters the duodenum, the bile is secreted from the gallbladder into the duodenum.
Summary
Different animals have evolved different types of digestive systems specialized to meet their dietary needs. Humans and many other animals have monogastric digestive systems with a single-chambered stomach. Birds have evolved a digestive system that includes a gizzard where the food is crushed into smaller pieces. This compensates for their inability to masticate. Ruminants that consume large amounts of plant material have a multi-chambered stomach that digests roughage. Pseudo-ruminants have similar digestive processes as ruminants but do not have the four-compartment stomach. Processing food involves ingestion (eating), digestion (mechanical and enzymatic breakdown of large molecules), absorption (cellular uptake of nutrients), and elimination (removal of undigested waste as feces).
Many organs work together to digest food and absorb nutrients. The mouth is the point of ingestion and the location where both mechanical and chemical breakdown of food begins. Saliva contains an enzyme called amylase that breaks down carbohydrates. The food bolus travels through the esophagus by peristaltic movements to the stomach. The stomach has an extremely acidic environment. An enzyme called pepsin digests protein in the stomach. Further digestion and absorption take place in the small intestine. The large intestine reabsorbs water from the undigested food and stores waste until elimination.
Art Connections
Figure \(10\): Which of the following statements about the digestive system is false?
1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.
Answer
B
Figure \(11\): Which of the following statements about the small intestine is false?
1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.
Answer
C
Glossary
alimentary canal
tubular digestive system with a mouth and anus
anus
exit point for waste material
bile
digestive juice produced by the liver; important for digestion of lipids
bolus
mass of food resulting from chewing action and wetting by saliva
carnivore
animal that consumes animal flesh
chyme
mixture of partially digested food and stomach juices
duodenum
first part of the small intestine where a large part of digestion of carbohydrates and fats occurs
esophagus
tubular organ that connects the mouth to the stomach
gallbladder
organ that stores and concentrates bile
gastrovascular cavity
digestive system consisting of a single opening
gizzard
muscular organ that grinds food
herbivore
animal that consumes strictly plant diet
ileum
last part of the small intestine; connects the small intestine to the large intestine; important for absorption of B-12
jejunum
second part of the small intestine
large intestine
digestive system organ that reabsorbs water from undigested material and processes waste matter
lipase
enzyme that chemically breaks down lipids
liver
organ that produces bile for digestion and processes vitamins and lipids
monogastric
digestive system that consists of a single-chambered stomach
omnivore
animal that consumes both plants and animals
pancreas
gland that secretes digestive juices
pepsin
enzyme found in the stomach whose main role is protein digestion
pepsinogen
inactive form of pepsin
peristalsis
wave-like movements of muscle tissue
proventriculus
glandular part of a bird’s stomach
rectum
area of the body where feces is stored until elimination
roughage
component of food that is low in energy and high in fiber
ruminant
animal with a stomach divided into four compartments
salivary amylase
enzyme found in saliva, which converts carbohydrates to maltose
small intestine
organ where digestion of protein, fats, and carbohydrates is completed
sphincter
band of muscle that controls movement of materials throughout the digestive tract
stomach
saclike organ containing acidic digestive juices
villi
folds on the inner surface of the small intestine whose role is to increase absorption area | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/46%3A_The_Digestive_System/46.02%3A_The_Mouth_and_Teeth-_Food_Capture_and_Bulk_Processing.txt |
Skills to Develop
• Describe the process of digestion
• Detail the steps involved in digestion and absorption
• Define elimination
• Explain the role of both the small and large intestines in absorption
Obtaining nutrition and energy from food is a multi-step process. For true animals, the first step is ingestion, the act of taking in food. This is followed by digestion, absorption, and elimination. In the following sections, each of these steps will be discussed in detail.
Ingestion
The large molecules found in intact food cannot pass through the cell membranes. Food needs to be broken into smaller particles so that animals can harness the nutrients and organic molecules. The first step in this process is ingestion. Ingestion is the process of taking in food through the mouth. In vertebrates, the teeth, saliva, and tongue play important roles in mastication (preparing the food into bolus). While the food is being mechanically broken down, the enzymes in saliva begin to chemically process the food as well. The combined action of these processes modifies the food from large particles to a soft mass that can be swallowed and can travel the length of the esophagus.
Digestion and Absorption
Digestion is the mechanical and chemical break down of food into small organic fragments. It is important to break down macromolecules into smaller fragments that are of suitable size for absorption across the digestive epithelium. Large, complex molecules of proteins, polysaccharides, and lipids must be reduced to simpler particles such as simple sugar before they can be absorbed by the digestive epithelial cells. Different organs play specific roles in the digestive process. The animal diet needs carbohydrates, protein, and fat, as well as vitamins and inorganic components for nutritional balance. How each of these components is digested is discussed in the following sections.
Carbohydrates
The digestion of carbohydrates begins in the mouth. The salivary enzyme amylase begins the breakdown of food starches into maltose, a disaccharide. As the bolus of food travels through the esophagus to the stomach, no significant digestion of carbohydrates takes place. The esophagus produces no digestive enzymes but does produce mucous for lubrication. The acidic environment in the stomach stops the action of the amylase enzyme.
The next step of carbohydrate digestion takes place in the duodenum. Recall that the chyme from the stomach enters the duodenum and mixes with the digestive secretion from the pancreas, liver, and gallbladder. Pancreatic juices also contain amylase, which continues the breakdown of starch and glycogen into maltose, a disaccharide. The disaccharides are broken down into monosaccharides by enzymes called maltases, sucrases, and lactases, which are also present in the brush border of the small intestinal wall. Maltase breaks down maltose into glucose. Other disaccharides, such as sucrose and lactose are broken down by sucrase and lactase, respectively. Sucrase breaks down sucrose (or “table sugar”) into glucose and fructose, and lactase breaks down lactose (or “milk sugar”) into glucose and galactose. The monosaccharides (glucose) thus produced are absorbed and then can be used in metabolic pathways to harness energy. The monosaccharides are transported across the intestinal epithelium into the bloodstream to be transported to the different cells in the body. The steps in carbohydrate digestion are summarized in Figure \(1\).
Table \(1\): Digestion of Carbohydrates
Enzyme Produced By Site of Action Substrate Acting On End Products
Salivary amylase Salivary glands Mouth Polysaccharides (Starch) Disaccharides (maltose), oligosaccharides
Pancreatic amylase Pancreas Small intestine Polysaccharides (starch) Disaccharides (maltose), monosaccharides
Oligosaccharidases Lining of the intestine; brush border membrane Small intestine Disaccharides Monosaccharides (e.g., glucose, fructose, galactose)
Protein
A large part of protein digestion takes place in the stomach. The enzyme pepsin plays an important role in the digestion of proteins by breaking down the intact protein to peptides, which are short chains of four to nine amino acids. In the duodenum, other enzymes—trypsin, elastase, and chymotrypsin—act on the peptides reducing them to smaller peptides. Trypsin elastase, carboxypeptidase, and chymotrypsin are produced by the pancreas and released into the duodenum where they act on the chyme. Further breakdown of peptides to single amino acids is aided by enzymes called peptidases (those that break down peptides). Specifically, carboxypeptidase, dipeptidase, and aminopeptidase play important roles in reducing the peptides to free amino acids. The amino acids are absorbed into the bloodstream through the small intestines. The steps in protein digestion are summarized in Figure \(2\).
Table \(2\): Digestion of Protein
Enzyme Produced By Site of Action Substrate Acting On End Products
Pepsin Stomach chief cells Stomach Proteins Peptides
Trypsin
Elastase Chymotrypsin
Pancreas Small intestine Proteins Peptides
Carboxypeptidase Pancreas Small intestine Peptides Amino acids and peptides
Aminopeptidase
Dipeptidase
Lining of intestine Small intestine Peptides Amino acids
Lipids
Lipid digestion begins in the stomach with the aid of lingual lipase and gastric lipase. However, the bulk of lipid digestion occurs in the small intestine due to pancreatic lipase. When chyme enters the duodenum, the hormonal responses trigger the release of bile, which is produced in the liver and stored in the gallbladder. Bile aids in the digestion of lipids, primarily triglycerides by emulsification. Emulsification is a process in which large lipid globules are broken down into several small lipid globules. These small globules are more widely distributed in the chyme rather than forming large aggregates. Lipids are hydrophobic substances: in the presence of water, they will aggregate to form globules to minimize exposure to water. Bile contains bile salts, which are amphipathic, meaning they contain hydrophobic and hydrophilic parts. Thus, the bile salts hydrophilic side can interface with water on one side and the hydrophobic side interfaces with lipids on the other. By doing so, bile salts emulsify large lipid globules into small lipid globules.
Why is emulsification important for digestion of lipids? Pancreatic juices contain enzymes called lipases (enzymes that break down lipids). If the lipid in the chyme aggregates into large globules, very little surface area of the lipids is available for the lipases to act on, leaving lipid digestion incomplete. By forming an emulsion, bile salts increase the available surface area of the lipids many fold. The pancreatic lipases can then act on the lipids more efficiently and digest them, as detailed in Figure \(3\). Lipases break down the lipids into fatty acids and glycerides. These molecules can pass through the plasma membrane of the cell and enter the epithelial cells of the intestinal lining. The bile salts surround long-chain fatty acids and monoglycerides forming tiny spheres called micelles. The micelles move into the brush border of the small intestine absorptive cells where the long-chain fatty acids and monoglycerides diffuse out of the micelles into the absorptive cells leaving the micelles behind in the chyme. The long-chain fatty acids and monoglycerides recombine in the absorptive cells to form triglycerides, which aggregate into globules and become coated with proteins. These large spheres are called chylomicrons. Chylomicrons contain triglycerides, cholesterol, and other lipids and have proteins on their surface. The surface is also composed of the hydrophilic phosphate "heads" of phospholipids. Together, they enable the chylomicron to move in an aqueous environment without exposing the lipids to water. Chylomicrons leave the absorptive cells via exocytosis. Chylomicrons enter the lymphatic vessels, and then enter the blood in the subclavian vein.
Vitamins
Vitamins can be either water-soluble or lipid-soluble. Fat soluble vitamins are absorbed in the same manner as lipids. It is important to consume some amount of dietary lipid to aid the absorption of lipid-soluble vitamins. Water-soluble vitamins can be directly absorbed into the bloodstream from the intestine.
Art Connection
Which of the following statements about digestive processes is true?
1. Amylase, maltase, and lactase in the mouth digest carbohydrates.
2. Trypsin and lipase in the stomach digest protein.
3. Bile emulsifies lipids in the small intestine.
4. No food is absorbed until the small intestine.
Elimination
The final step in digestion is the elimination of undigested food content and waste products. The undigested food material enters the colon, where most of the water is reabsorbed. Recall that the colon is also home to the microflora called “intestinal flora” that aid in the digestion process. The semi-solid waste is moved through the colon by peristaltic movements of the muscle and is stored in the rectum. As the rectum expands in response to storage of fecal matter, it triggers the neural signals required to set up the urge to eliminate. The solid waste is eliminated through the anus using peristaltic movements of the rectum.
Common Problems with Elimination
Diarrhea and constipation are some of the most common health concerns that affect digestion. Constipation is a condition where the feces are hardened because of excess water removal in the colon. In contrast, if enough water is not removed from the feces, it results in diarrhea. Many bacteria, including the ones that cause cholera, affect the proteins involved in water reabsorption in the colon and result in excessive diarrhea.
Emesis
Emesis, or vomiting, is elimination of food by forceful expulsion through the mouth. It is often in response to an irritant that affects the digestive tract, including but not limited to viruses, bacteria, emotions, sights, and food poisoning. This forceful expulsion of the food is due to the strong contractions produced by the stomach muscles. The process of emesis is regulated by the medulla.
Summary
Digestion begins with ingestion, where the food is taken in the mouth. Digestion and absorption take place in a series of steps with special enzymes playing important roles in digesting carbohydrates, proteins, and lipids. Elimination describes removal of undigested food contents and waste products from the body. While most absorption occurs in the small intestines, the large intestine is responsible for the final removal of water that remains after the absorptive process of the small intestines. The cells that line the large intestine absorb some vitamins as well as any leftover salts and water. The large intestine (colon) is also where feces is formed.
Art Connections
Figure \(4\): Which of the following statements about digestive processes is true?
1. Amylase, maltase and lactase in the mouth digest carbohydrates.
2. Trypsin and lipase in the stomach digest protein.
3. Bile emulsifies lipids in the small intestine.
4. No food is absorbed until the small intestine.
Answer
C
Glossary
aminopeptidase
protease that breaks down peptides to single amino acids; secreted by the brush border of small intestine
carboxypeptidase
protease that breaks down peptides to single amino acids; secreted by the brush border of the small intestine
chylomicron
small lipid globule
chymotrypsin
pancreatic protease
digestion
mechanical and chemical break down of food into small organic fragments
dipeptidase
protease that breaks down peptides to single amino acids; secreted by the brush border of small intestine
elastase
pancreatic protease
ingestion
act of taking in food
lactase
enzyme that breaks down lactose into glucose and galactose
maltase
enzyme that breaks down maltose into glucose
sucrase
enzyme that breaks down sucrose into glucose and fructose
trypsin
pancreatic protease that breaks down protein | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/46%3A_The_Digestive_System/46.03%3A_The_Esophagus_and_Stomach-_The_Early_Stages_of_Digestion.txt |
Skills to Develop
• Describe the process of digestion
• Detail the steps involved in digestion and absorption
• Define elimination
• Explain the role of both the small and large intestines in absorption
Obtaining nutrition and energy from food is a multi-step process. For true animals, the first step is ingestion, the act of taking in food. This is followed by digestion, absorption, and elimination. In the following sections, each of these steps will be discussed in detail.
Ingestion
The large molecules found in intact food cannot pass through the cell membranes. Food needs to be broken into smaller particles so that animals can harness the nutrients and organic molecules. The first step in this process is ingestion. Ingestion is the process of taking in food through the mouth. In vertebrates, the teeth, saliva, and tongue play important roles in mastication (preparing the food into bolus). While the food is being mechanically broken down, the enzymes in saliva begin to chemically process the food as well. The combined action of these processes modifies the food from large particles to a soft mass that can be swallowed and can travel the length of the esophagus.
Digestion and Absorption
Digestion is the mechanical and chemical break down of food into small organic fragments. It is important to break down macromolecules into smaller fragments that are of suitable size for absorption across the digestive epithelium. Large, complex molecules of proteins, polysaccharides, and lipids must be reduced to simpler particles such as simple sugar before they can be absorbed by the digestive epithelial cells. Different organs play specific roles in the digestive process. The animal diet needs carbohydrates, protein, and fat, as well as vitamins and inorganic components for nutritional balance. How each of these components is digested is discussed in the following sections.
Carbohydrates
The digestion of carbohydrates begins in the mouth. The salivary enzyme amylase begins the breakdown of food starches into maltose, a disaccharide. As the bolus of food travels through the esophagus to the stomach, no significant digestion of carbohydrates takes place. The esophagus produces no digestive enzymes but does produce mucous for lubrication. The acidic environment in the stomach stops the action of the amylase enzyme.
The next step of carbohydrate digestion takes place in the duodenum. Recall that the chyme from the stomach enters the duodenum and mixes with the digestive secretion from the pancreas, liver, and gallbladder. Pancreatic juices also contain amylase, which continues the breakdown of starch and glycogen into maltose, a disaccharide. The disaccharides are broken down into monosaccharides by enzymes called maltases, sucrases, and lactases, which are also present in the brush border of the small intestinal wall. Maltase breaks down maltose into glucose. Other disaccharides, such as sucrose and lactose are broken down by sucrase and lactase, respectively. Sucrase breaks down sucrose (or “table sugar”) into glucose and fructose, and lactase breaks down lactose (or “milk sugar”) into glucose and galactose. The monosaccharides (glucose) thus produced are absorbed and then can be used in metabolic pathways to harness energy. The monosaccharides are transported across the intestinal epithelium into the bloodstream to be transported to the different cells in the body. The steps in carbohydrate digestion are summarized in Figure \(1\).
Table \(1\): Digestion of Carbohydrates
Enzyme Produced By Site of Action Substrate Acting On End Products
Salivary amylase Salivary glands Mouth Polysaccharides (Starch) Disaccharides (maltose), oligosaccharides
Pancreatic amylase Pancreas Small intestine Polysaccharides (starch) Disaccharides (maltose), monosaccharides
Oligosaccharidases Lining of the intestine; brush border membrane Small intestine Disaccharides Monosaccharides (e.g., glucose, fructose, galactose)
Protein
A large part of protein digestion takes place in the stomach. The enzyme pepsin plays an important role in the digestion of proteins by breaking down the intact protein to peptides, which are short chains of four to nine amino acids. In the duodenum, other enzymes—trypsin, elastase, and chymotrypsin—act on the peptides reducing them to smaller peptides. Trypsin elastase, carboxypeptidase, and chymotrypsin are produced by the pancreas and released into the duodenum where they act on the chyme. Further breakdown of peptides to single amino acids is aided by enzymes called peptidases (those that break down peptides). Specifically, carboxypeptidase, dipeptidase, and aminopeptidase play important roles in reducing the peptides to free amino acids. The amino acids are absorbed into the bloodstream through the small intestines. The steps in protein digestion are summarized in Figure \(2\).
Table \(2\): Digestion of Protein
Enzyme Produced By Site of Action Substrate Acting On End Products
Pepsin Stomach chief cells Stomach Proteins Peptides
Trypsin
Elastase Chymotrypsin
Pancreas Small intestine Proteins Peptides
Carboxypeptidase Pancreas Small intestine Peptides Amino acids and peptides
Aminopeptidase
Dipeptidase
Lining of intestine Small intestine Peptides Amino acids
Lipids
Lipid digestion begins in the stomach with the aid of lingual lipase and gastric lipase. However, the bulk of lipid digestion occurs in the small intestine due to pancreatic lipase. When chyme enters the duodenum, the hormonal responses trigger the release of bile, which is produced in the liver and stored in the gallbladder. Bile aids in the digestion of lipids, primarily triglycerides by emulsification. Emulsification is a process in which large lipid globules are broken down into several small lipid globules. These small globules are more widely distributed in the chyme rather than forming large aggregates. Lipids are hydrophobic substances: in the presence of water, they will aggregate to form globules to minimize exposure to water. Bile contains bile salts, which are amphipathic, meaning they contain hydrophobic and hydrophilic parts. Thus, the bile salts hydrophilic side can interface with water on one side and the hydrophobic side interfaces with lipids on the other. By doing so, bile salts emulsify large lipid globules into small lipid globules.
Why is emulsification important for digestion of lipids? Pancreatic juices contain enzymes called lipases (enzymes that break down lipids). If the lipid in the chyme aggregates into large globules, very little surface area of the lipids is available for the lipases to act on, leaving lipid digestion incomplete. By forming an emulsion, bile salts increase the available surface area of the lipids many fold. The pancreatic lipases can then act on the lipids more efficiently and digest them, as detailed in Figure \(3\). Lipases break down the lipids into fatty acids and glycerides. These molecules can pass through the plasma membrane of the cell and enter the epithelial cells of the intestinal lining. The bile salts surround long-chain fatty acids and monoglycerides forming tiny spheres called micelles. The micelles move into the brush border of the small intestine absorptive cells where the long-chain fatty acids and monoglycerides diffuse out of the micelles into the absorptive cells leaving the micelles behind in the chyme. The long-chain fatty acids and monoglycerides recombine in the absorptive cells to form triglycerides, which aggregate into globules and become coated with proteins. These large spheres are called chylomicrons. Chylomicrons contain triglycerides, cholesterol, and other lipids and have proteins on their surface. The surface is also composed of the hydrophilic phosphate "heads" of phospholipids. Together, they enable the chylomicron to move in an aqueous environment without exposing the lipids to water. Chylomicrons leave the absorptive cells via exocytosis. Chylomicrons enter the lymphatic vessels, and then enter the blood in the subclavian vein.
Vitamins
Vitamins can be either water-soluble or lipid-soluble. Fat soluble vitamins are absorbed in the same manner as lipids. It is important to consume some amount of dietary lipid to aid the absorption of lipid-soluble vitamins. Water-soluble vitamins can be directly absorbed into the bloodstream from the intestine.
Art Connection
Which of the following statements about digestive processes is true?
1. Amylase, maltase, and lactase in the mouth digest carbohydrates.
2. Trypsin and lipase in the stomach digest protein.
3. Bile emulsifies lipids in the small intestine.
4. No food is absorbed until the small intestine.
Elimination
The final step in digestion is the elimination of undigested food content and waste products. The undigested food material enters the colon, where most of the water is reabsorbed. Recall that the colon is also home to the microflora called “intestinal flora” that aid in the digestion process. The semi-solid waste is moved through the colon by peristaltic movements of the muscle and is stored in the rectum. As the rectum expands in response to storage of fecal matter, it triggers the neural signals required to set up the urge to eliminate. The solid waste is eliminated through the anus using peristaltic movements of the rectum.
Common Problems with Elimination
Diarrhea and constipation are some of the most common health concerns that affect digestion. Constipation is a condition where the feces are hardened because of excess water removal in the colon. In contrast, if enough water is not removed from the feces, it results in diarrhea. Many bacteria, including the ones that cause cholera, affect the proteins involved in water reabsorption in the colon and result in excessive diarrhea.
Emesis
Emesis, or vomiting, is elimination of food by forceful expulsion through the mouth. It is often in response to an irritant that affects the digestive tract, including but not limited to viruses, bacteria, emotions, sights, and food poisoning. This forceful expulsion of the food is due to the strong contractions produced by the stomach muscles. The process of emesis is regulated by the medulla.
Summary
Digestion begins with ingestion, where the food is taken in the mouth. Digestion and absorption take place in a series of steps with special enzymes playing important roles in digesting carbohydrates, proteins, and lipids. Elimination describes removal of undigested food contents and waste products from the body. While most absorption occurs in the small intestines, the large intestine is responsible for the final removal of water that remains after the absorptive process of the small intestines. The cells that line the large intestine absorb some vitamins as well as any leftover salts and water. The large intestine (colon) is also where feces is formed.
Art Connections
Figure \(4\): Which of the following statements about digestive processes is true?
1. Amylase, maltase and lactase in the mouth digest carbohydrates.
2. Trypsin and lipase in the stomach digest protein.
3. Bile emulsifies lipids in the small intestine.
4. No food is absorbed until the small intestine.
Answer
C
Glossary
aminopeptidase
protease that breaks down peptides to single amino acids; secreted by the brush border of small intestine
carboxypeptidase
protease that breaks down peptides to single amino acids; secreted by the brush border of the small intestine
chylomicron
small lipid globule
chymotrypsin
pancreatic protease
digestion
mechanical and chemical break down of food into small organic fragments
dipeptidase
protease that breaks down peptides to single amino acids; secreted by the brush border of small intestine
elastase
pancreatic protease
ingestion
act of taking in food
lactase
enzyme that breaks down lactose into glucose and galactose
maltase
enzyme that breaks down maltose into glucose
sucrase
enzyme that breaks down sucrose into glucose and fructose
trypsin
pancreatic protease that breaks down protein | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/46%3A_The_Digestive_System/46.04%3A_The_Intestines-_Breakdown_Absorption_and_Elimination.txt |
Skills to Develop
• Discuss the role of neural regulation in digestive processes
• Explain how hormones regulate digestion
The brain is the control center for the sensation of hunger and satiety. The functions of the digestive system are regulated through neural and hormonal responses.
Neural Responses to Food
In reaction to the smell, sight, or thought of food, like that shown in Figure \(1\), the first hormonal response is that of salivation. The salivary glands secrete more saliva in response to the stimulus presented by food in preparation for digestion. Simultaneously, the stomach begins to produce hydrochloric acid to digest the food. Recall that the peristaltic movements of the esophagus and other organs of the digestive tract are under the control of the brain. The brain prepares these muscles for movement as well. When the stomach is full, the part of the brain that detects satiety signals fullness. There are three overlapping phases of gastric control—the cephalic phase, the gastric phase, and the intestinal phase—each requires many enzymes and is under neural control as well.
Digestive Phases
The response to food begins even before food enters the mouth. The first phase of ingestion, called the cephalic phase, is controlled by the neural response to the stimulus provided by food. All aspects—such as sight, sense, and smell—trigger the neural responses resulting in salivation and secretion of gastric juices. The gastric and salivary secretion in the cephalic phase can also take place due to the thought of food. Right now, if you think about a piece of chocolate or a crispy potato chip, the increase in salivation is a cephalic phase response to the thought. The central nervous system prepares the stomach to receive food.
The gastric phase begins once the food arrives in the stomach. It builds on the stimulation provided during the cephalic phase. Gastric acids and enzymes process the ingested materials. The gastric phase is stimulated by (1) distension of the stomach, (2) a decrease in the pH of the gastric contents, and (3) the presence of undigested material. This phase consists of local, hormonal, and neural responses. These responses stimulate secretions and powerful contractions.
The intestinal phase begins when chyme enters the small intestine triggering digestive secretions. This phase controls the rate of gastric emptying. In addition to gastrin emptying, when chyme enters the small intestine, it triggers other hormonal and neural events that coordinate the activities of the intestinal tract, pancreas, liver, and gallbladder.
Hormonal Responses to Food
The endocrine system controls the response of the various glands in the body and the release of hormones at the appropriate times.
One of the important factors under hormonal control is the stomach acid environment. During the gastric phase, the hormone gastrin is secreted by G cells in the stomach in response to the presence of proteins. Gastrin stimulates the release of stomach acid, or hydrochloric acid (HCl) which aids in the digestion of the proteins. However, when the stomach is emptied, the acidic environment need not be maintained and a hormone called somatostatin stops the release of hydrochloric acid. This is controlled by a negative feedback mechanism.
In the duodenum, digestive secretions from the liver, pancreas, and gallbladder play an important role in digesting chyme during the intestinal phase. In order to neutralize the acidic chyme, a hormone called secretin stimulates the pancreas to produce alkaline bicarbonate solution and deliver it to the duodenum. Secretin acts in tandem with another hormone called cholecystokinin (CCK). Not only does CCK stimulate the pancreas to produce the requisite pancreatic juices, it also stimulates the gallbladder to release bile into the duodenum.
Another level of hormonal control occurs in response to the composition of food. Foods high in lipids take a long time to digest. A hormone called gastric inhibitory peptide is secreted by the small intestine to slow down the peristaltic movements of the intestine to allow fatty foods more time to be digested and absorbed.
Understanding the hormonal control of the digestive system is an important area of ongoing research. Scientists are exploring the role of each hormone in the digestive process and developing ways to target these hormones. Advances could lead to knowledge that may help to battle the obesity epidemic.
Summary
The brain and the endocrine system control digestive processes. The brain controls the responses of hunger and satiety. The endocrine system controls the release of hormones and enzymes required for digestion of food in the digestive tract.
Glossary
cephalic phase
first phase of digestion, controlled by the neural response to the stimulus provided by food
cholecystokinin
hormone that stimulates the contraction of the gallbladder to release bile
endocrine system
system that controls the response of the various glands in the body and the release of hormones at the appropriate times
gastric inhibitory peptide
hormone secreted by the small intestine in the presence of fatty acids and sugars; it also inhibits acid production and peristalsis in order to slow down the rate at which food enters the small intestine
gastric phase
digestive phase beginning once food enters the stomach; gastric acids and enzymes process the ingested materials
gastrin
hormone which stimulates hydrochloric acid secretion in the stomach
intestinal phase
third digestive phase; begins when chyme enters the small intestine triggering digestive secretions and controlling the rate of gastric emptying
secretin
hormone which stimulates sodium bicarbonate secretion in the small intestine
somatostatin
hormone released to stop acid secretion when the stomach is empty | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/46%3A_The_Digestive_System/46.06%3A_Neural_and_Hormonal_Regulation_of_the_Digestive_Tract.txt |
Skills to Develop
• Explain the processes of digestion and absorption
• Compare and contrast different types of digestive systems
• Explain the specialized functions of the organs involved in processing food in the body
• Describe the ways in which organs work together to digest food and absorb nutrients
Animals obtain their nutrition from the consumption of other organisms. Depending on their diet, animals can be classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not immediately accessible to the cells. There are a number of processes that modify food within the animal body in order to make the nutrients and organic molecules accessible for cellular function. As animals evolved in complexity of form and function, their digestive systems have also evolved to accommodate their various dietary needs.
Herbivores, Omnivores, and Carnivores
Herbivores are animals whose primary food source is plant-based. Examples of herbivores, as shown in Figure \(1\) include vertebrates like deer, koalas, and some bird species, as well as invertebrates such as crickets and caterpillars. These animals have evolved digestive systems capable of handling large amounts of plant material. Herbivores can be further classified into frugivores (fruit-eaters), granivores (seed eaters), nectivores (nectar feeders), and folivores (leaf eaters).
Carnivores are animals that eat other animals. The word carnivore is derived from Latin and literally means “meat eater.” Wild cats such as lions, shown in Figure \(2\)a and tigers are examples of vertebrate carnivores, as are snakes and sharks, while invertebrate carnivores include sea stars, spiders, and ladybugs, shown in Figure \(2\)b. Obligate carnivores are those that rely entirely on animal flesh to obtain their nutrients; examples of obligate carnivores are members of the cat family, such as lions and cheetahs. Facultative carnivores are those that also eat non-animal food in addition to animal food. Note that there is no clear line that differentiates facultative carnivores from omnivores; dogs would be considered facultative carnivores.
Omnivores are animals that eat both plant- and animal-derived food. In Latin, omnivore means to eat everything. Humans, bears (shown in Figure \(3\)a), and chickens are example of vertebrate omnivores; invertebrate omnivores include cockroaches and crayfish (shown in Figure \(3\)b).
Invertebrate Digestive Systems
Animals have evolved different types of digestive systems to aid in the digestion of the different foods they consume. The simplest example is that of a gastrovascular cavity and is found in organisms with only one opening for digestion. Platyhelminthes (flatworms), Ctenophora (comb jellies), and Cnidaria (coral, jelly fish, and sea anemones) use this type of digestion. Gastrovascular cavities, as shown in Figure \(4\)a, are typically a blind tube or cavity with only one opening, the “mouth”, which also serves as an “anus”. Ingested material enters the mouth and passes through a hollow, tubular cavity. Cells within the cavity secrete digestive enzymes that break down the food. The food particles are engulfed by the cells lining the gastrovascular cavity.
The alimentary canal, shown in Figure \(4\)b, is a more advanced system: it consists of one tube with a mouth at one end and an anus at the other. Earthworms are an example of an animal with an alimentary canal. Once the food is ingested through the mouth, it passes through the esophagus and is stored in an organ called the crop; then it passes into the gizzard where it is churned and digested. From the gizzard, the food passes through the intestine, the nutrients are absorbed, and the waste is eliminated as feces, called castings, through the anus.
Vertebrate Digestive Systems
Vertebrates have evolved more complex digestive systems to adapt to their dietary needs. Some animals have a single stomach, while others have multi-chambered stomachs. Birds have developed a digestive system adapted to eating unmasticated food.
Monogastric: Single-chambered Stomach
As the word monogastric suggests, this type of digestive system consists of one (“mono”) stomach chamber (“gastric”). Humans and many animals have a monogastric digestive system as illustrated in Figure \(5\). The process of digestion begins with the mouth and the intake of food. The teeth play an important role in masticating (chewing) or physically breaking down food into smaller particles. The enzymes present in saliva also begin to chemically break down food. The esophagus is a long tube that connects the mouth to the stomach. Using peristalsis, or wave-like smooth muscle contractions, the muscles of the esophagus push the food towards the stomach. In order to speed up the actions of enzymes in the stomach, the stomach is an extremely acidic environment, with a pH between 1.5 and 2.5. The gastric juices, which include enzymes in the stomach, act on the food particles and continue the process of digestion. Further breakdown of food takes place in the small intestine where enzymes produced by the liver, the small intestine, and the pancreas continue the process of digestion. The nutrients are absorbed into the blood stream across the epithelial cells lining the walls of the small intestines. The waste material travels on to the large intestine where water is absorbed and the drier waste material is compacted into feces; it is stored until it is excreted through the rectum.
Avian
Birds face special challenges when it comes to obtaining nutrition from food. They do not have teeth and so their digestive system, shown in Figure \(6\), must be able to process un-masticated food. Birds have evolved a variety of beak types that reflect the vast variety in their diet, ranging from seeds and insects to fruits and nuts. Because most birds fly, their metabolic rates are high in order to efficiently process food and keep their body weight low. The stomach of birds has two chambers: the proventriculus, where gastric juices are produced to digest the food before it enters the stomach, and the gizzard, where the food is stored, soaked, and mechanically ground. The undigested material forms food pellets that are sometimes regurgitated. Most of the chemical digestion and absorption happens in the intestine and the waste is excreted through the cloaca.
Evolution Connection: Avian Adaptations
Birds have a highly efficient, simplified digestive system. Recent fossil evidence has shown that the evolutionary divergence of birds from other land animals was characterized by streamlining and simplifying the digestive system. Unlike many other animals, birds do not have teeth to chew their food. In place of lips, they have sharp pointy beaks. The horny beak, lack of jaws, and the smaller tongue of the birds can be traced back to their dinosaur ancestors. The emergence of these changes seems to coincide with the inclusion of seeds in the bird diet. Seed-eating birds have beaks that are shaped for grabbing seeds and the two-compartment stomach allows for delegation of tasks. Since birds need to remain light in order to fly, their metabolic rates are very high, which means they digest their food very quickly and need to eat often. Contrast this with the ruminants, where the digestion of plant matter takes a very long time.
Ruminants
Ruminants are mainly herbivores like cows, sheep, and goats, whose entire diet consists of eating large amounts of roughage or fiber. They have evolved digestive systems that help them digest vast amounts of cellulose. An interesting feature of the ruminants’ mouth is that they do not have upper incisor teeth. They use their lower teeth, tongue and lips to tear and chew their food. From the mouth, the food travels to the esophagus and on to the stomach.
To help digest the large amount of plant material, the stomach of the ruminants is a multi-chambered organ, as illustrated in Figure \(7\). The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that break down cellulose and ferment ingested food. The abomasum is the “true” stomach and is the equivalent of the monogastric stomach chamber where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the microbial support necessary to digest plant material in ruminants. The fermentation process produces large amounts of gas in the stomach chamber, which must be eliminated. As in other animals, the small intestine plays an important role in nutrient absorption, and the large intestine helps in the elimination of waste.
Pseudo-ruminants
Some animals, such as camels and alpacas, are pseudo-ruminants. They eat a lot of plant material and roughage. Digesting plant material is not easy because plant cell walls contain the polymeric sugar molecule cellulose. The digestive enzymes of these animals cannot break down cellulose, but microorganisms present in the digestive system can. Therefore, the digestive system must be able to handle large amounts of roughage and break down the cellulose. Pseudo-ruminants have a three-chamber stomach in the digestive system. However, their cecum—a pouched organ at the beginning of the large intestine containing many microorganisms that are necessary for the digestion of plant materials—is large and is the site where the roughage is fermented and digested. These animals do not have a rumen but have an omasum, abomasum, and reticulum.
Parts of the Digestive System
The vertebrate digestive system is designed to facilitate the transformation of food matter into the nutrient components that sustain organisms.
Oral Cavity
The oral cavity, or mouth, is the point of entry of food into the digestive system, illustrated in Figure \(8\). The food consumed is broken into smaller particles by mastication, the chewing action of the teeth. All mammals have teeth and can chew their food.
The extensive chemical process of digestion begins in the mouth. As food is being chewed, saliva, produced by the salivary glands, mixes with the food. Saliva is a watery substance produced in the mouths of many animals. There are three major glands that secrete saliva—the parotid, the submandibular, and the sublingual. Saliva contains mucus that moistens food and buffers the pH of the food. Saliva also contains immunoglobulins and lysozymes, which have antibacterial action to reduce tooth decay by inhibiting growth of some bacteria. Saliva also contains an enzyme called salivary amylase that begins the process of converting starches in the food into a disaccharide called maltose. Another enzyme called lipase is produced by the cells in the tongue. Lipases are a class of enzymes that can break down triglycerides. The lingual lipase begins the breakdown of fat components in the food. The chewing and wetting action provided by the teeth and saliva prepare the food into a mass called the bolus for swallowing. The tongue helps in swallowing—moving the bolus from the mouth into the pharynx. The pharynx opens to two passageways: the trachea, which leads to the lungs, and the esophagus, which leads to the stomach. The trachea has an opening called the glottis, which is covered by a cartilaginous flap called the epiglottis. When swallowing, the epiglottis closes the glottis and food passes into the esophagus and not the trachea. This arrangement allows food to be kept out of the trachea.
Esophagus
The esophagus is a tubular organ that connects the mouth to the stomach. The chewed and softened food passes through the esophagus after being swallowed. The smooth muscles of the esophagus undergo a series of wave like movements called peristalsis that push the food toward the stomach, as illustrated in Figure \(9\). The peristalsis wave is unidirectional—it moves food from the mouth to the stomach, and reverse movement is not possible. The peristaltic movement of the esophagus is an involuntary reflex; it takes place in response to the act of swallowing.
A ring-like muscle called a sphincter forms valves in the digestive system. The gastro-esophageal sphincter is located at the stomach end of the esophagus. In response to swallowing and the pressure exerted by the bolus of food, this sphincter opens, and the bolus enters the stomach. When there is no swallowing action, this sphincter is shut and prevents the contents of the stomach from traveling up the esophagus. Many animals have a true sphincter; however, in humans, there is no true sphincter, but the esophagus remains closed when there is no swallowing action. Acid reflux or “heartburn” occurs when the acidic digestive juices escape into the esophagus.
Stomach
A large part of digestion occurs in the stomach, shown in Figure \(10\). The stomach is a saclike organ that secretes gastric digestive juices. The pH in the stomach is between 1.5 and 2.5. This highly acidic environment is required for the chemical breakdown of food and the extraction of nutrients. When empty, the stomach is a rather small organ; however, it can expand to up to 20 times its resting size when filled with food. This characteristic is particularly useful for animals that need to eat when food is available.
Art Connection
Which of the following statements about the digestive system is false?
1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.
The stomach is also the major site for protein digestion in animals other than ruminants. Protein digestion is mediated by an enzyme called pepsin in the stomach chamber. Pepsin is secreted by the chief cells in the stomach in an inactive form called pepsinogen. Pepsin breaks peptide bonds and cleaves proteins into smaller polypeptides; it also helps activate more pepsinogen, starting a positive feedback mechanism that generates more pepsin. Another cell type—parietal cells—secrete hydrogen and chloride ions, which combine in the lumen to form hydrochloric acid, the primary acidic component of the stomach juices. Hydrochloric acid helps to convert the inactive pepsinogen to pepsin. The highly acidic environment also kills many microorganisms in the food and, combined with the action of the enzyme pepsin, results in the hydrolysis of protein in the food. Chemical digestion is facilitated by the churning action of the stomach. Contraction and relaxation of smooth muscles mixes the stomach contents about every 20 minutes. The partially digested food and gastric juice mixture is called chyme. Chyme passes from the stomach to the small intestine. Further protein digestion takes place in the small intestine. Gastric emptying occurs within two to six hours after a meal. Only a small amount of chyme is released into the small intestine at a time. The movement of chyme from the stomach into the small intestine is regulated by the pyloric sphincter.
When digesting protein and some fats, the stomach lining must be protected from getting digested by pepsin. There are two points to consider when describing how the stomach lining is protected. First, as previously mentioned, the enzyme pepsin is synthesized in the inactive form. This protects the chief cells, because pepsinogen does not have the same enzyme functionality of pepsin. Second, the stomach has a thick mucus lining that protects the underlying tissue from the action of the digestive juices. When this mucus lining is ruptured, ulcers can form in the stomach. Ulcers are open wounds in or on an organ caused by bacteria (Helicobacter pylori) when the mucus lining is ruptured and fails to reform.
Small Intestine
Chyme moves from the stomach to the small intestine. The small intestine is the organ where the digestion of protein, fats, and carbohydrates is completed. The small intestine is a long tube-like organ with a highly folded surface containing finger-like projections called the villi. The apical surface of each villus has many microscopic projections called microvilli. These structures, illustrated in Figure \(11\), are lined with epithelial cells on the luminal side and allow for the nutrients to be absorbed from the digested food and absorbed into the blood stream on the other side. The villi and microvilli, with their many folds, increase the surface area of the intestine and increase absorption efficiency of the nutrients. Absorbed nutrients in the blood are carried into the hepatic portal vein, which leads to the liver. There, the liver regulates the distribution of nutrients to the rest of the body and removes toxic substances, including drugs, alcohol, and some pathogens.
Art Connection
Which of the following statements about the small intestine is false?
1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.
The human small intestine is over 6m long and is divided into three parts: the duodenum, the jejunum, and the ileum. The “C-shaped,” fixed part of the small intestine is called the duodenum and is shown in Figure \(11\). The duodenum is separated from the stomach by the pyloric sphincter which opens to allow chyme to move from the stomach to the duodenum. In the duodenum, chyme is mixed with pancreatic juices in an alkaline solution rich in bicarbonate that neutralizes the acidity of chyme and acts as a buffer. Pancreatic juices also contain several digestive enzymes. Digestive juices from the pancreas, liver, and gallbladder, as well as from gland cells of the intestinal wall itself, enter the duodenum. Bile is produced in the liver and stored and concentrated in the gallbladder. Bile contains bile salts which emulsify lipids while the pancreas produces enzymes that catabolize starches, disaccharides, proteins, and fats. These digestive juices break down the food particles in the chyme into glucose, triglycerides, and amino acids. Some chemical digestion of food takes place in the duodenum. Absorption of fatty acids also takes place in the duodenum.
The second part of the small intestine is called the jejunum, shown in Figure \(11\). Here, hydrolysis of nutrients is continued while most of the carbohydrates and amino acids are absorbed through the intestinal lining. The bulk of chemical digestion and nutrient absorption occurs in the jejunum.
The ileum, also illustrated in Figure \(11\) is the last part of the small intestine and here the bile salts and vitamins are absorbed into blood stream. The undigested food is sent to the colon from the ileum via peristaltic movements of the muscle. The ileum ends and the large intestine begins at the ileocecal valve. The vermiform, “worm-like,” appendix is located at the ileocecal valve. The appendix of humans secretes no enzymes and has an insignificant role in immunity.
Large Intestine
The large intestine, illustrated in Figure \(12\), reabsorbs the water from the undigested food material and processes the waste material. The human large intestine is much smaller in length compared to the small intestine but larger in diameter. It has three parts: the cecum, the colon, and the rectum. The cecum joins the ileum to the colon and is the receiving pouch for the waste matter. The colon is home to many bacteria or “intestinal flora” that aid in the digestive processes. The colon can be divided into four regions, the ascending colon, the transverse colon, the descending colon and the sigmoid colon. The main functions of the colon are to extract the water and mineral salts from undigested food, and to store waste material. Carnivorous mammals have a shorter large intestine compared to herbivorous mammals due to their diet.
Rectum and Anus
The rectum is the terminal end of the large intestine, as shown in Figure \(12\). The primary role of the rectum is to store the feces until defecation. The feces are propelled using peristaltic movements during elimination. The anus is an opening at the far-end of the digestive tract and is the exit point for the waste material. Two sphincters between the rectum and anus control elimination: the inner sphincter is involuntary and the outer sphincter is voluntary.
Accessory Organs
The organs discussed above are the organs of the digestive tract through which food passes. Accessory organs are organs that add secretions (enzymes) that catabolize food into nutrients. Accessory organs include salivary glands, the liver, the pancreas, and the gallbladder. The liver, pancreas, and gallbladder are regulated by hormones in response to the food consumed.
The liver is the largest internal organ in humans and it plays a very important role in digestion of fats and detoxifying blood. The liver produces bile, a digestive juice that is required for the breakdown of fatty components of the food in the duodenum. The liver also processes the vitamins and fats and synthesizes many plasma proteins.
The pancreas is another important gland that secretes digestive juices. The chyme produced from the stomach is highly acidic in nature; the pancreatic juices contain high levels of bicarbonate, an alkali that neutralizes the acidic chyme. Additionally, the pancreatic juices contain a large variety of enzymes that are required for the digestion of protein and carbohydrates.
The gallbladder is a small organ that aids the liver by storing bile and concentrating bile salts. When chyme containing fatty acids enters the duodenum, the bile is secreted from the gallbladder into the duodenum.
Summary
Different animals have evolved different types of digestive systems specialized to meet their dietary needs. Humans and many other animals have monogastric digestive systems with a single-chambered stomach. Birds have evolved a digestive system that includes a gizzard where the food is crushed into smaller pieces. This compensates for their inability to masticate. Ruminants that consume large amounts of plant material have a multi-chambered stomach that digests roughage. Pseudo-ruminants have similar digestive processes as ruminants but do not have the four-compartment stomach. Processing food involves ingestion (eating), digestion (mechanical and enzymatic breakdown of large molecules), absorption (cellular uptake of nutrients), and elimination (removal of undigested waste as feces).
Many organs work together to digest food and absorb nutrients. The mouth is the point of ingestion and the location where both mechanical and chemical breakdown of food begins. Saliva contains an enzyme called amylase that breaks down carbohydrates. The food bolus travels through the esophagus by peristaltic movements to the stomach. The stomach has an extremely acidic environment. An enzyme called pepsin digests protein in the stomach. Further digestion and absorption take place in the small intestine. The large intestine reabsorbs water from the undigested food and stores waste until elimination.
Art Connections
Figure \(10\): Which of the following statements about the digestive system is false?
1. Chyme is a mixture of food and digestive juices that is produced in the stomach.
2. Food enters the large intestine before the small intestine.
3. In the small intestine, chyme mixes with bile, which emulsifies fats.
4. The stomach is separated from the small intestine by the pyloric sphincter.
Answer
B
Figure \(11\): Which of the following statements about the small intestine is false?
1. Absorptive cells that line the small intestine have microvilli, small projections that increase surface area and aid in the absorption of food.
2. The inside of the small intestine has many folds, called villi.
3. Microvilli are lined with blood vessels as well as lymphatic vessels.
4. The inside of the small intestine is called the lumen.
Answer
C
Glossary
alimentary canal
tubular digestive system with a mouth and anus
anus
exit point for waste material
bile
digestive juice produced by the liver; important for digestion of lipids
bolus
mass of food resulting from chewing action and wetting by saliva
carnivore
animal that consumes animal flesh
chyme
mixture of partially digested food and stomach juices
duodenum
first part of the small intestine where a large part of digestion of carbohydrates and fats occurs
esophagus
tubular organ that connects the mouth to the stomach
gallbladder
organ that stores and concentrates bile
gastrovascular cavity
digestive system consisting of a single opening
gizzard
muscular organ that grinds food
herbivore
animal that consumes strictly plant diet
ileum
last part of the small intestine; connects the small intestine to the large intestine; important for absorption of B-12
jejunum
second part of the small intestine
large intestine
digestive system organ that reabsorbs water from undigested material and processes waste matter
lipase
enzyme that chemically breaks down lipids
liver
organ that produces bile for digestion and processes vitamins and lipids
monogastric
digestive system that consists of a single-chambered stomach
omnivore
animal that consumes both plants and animals
pancreas
gland that secretes digestive juices
pepsin
enzyme found in the stomach whose main role is protein digestion
pepsinogen
inactive form of pepsin
peristalsis
wave-like movements of muscle tissue
proventriculus
glandular part of a bird’s stomach
rectum
area of the body where feces is stored until elimination
roughage
component of food that is low in energy and high in fiber
ruminant
animal with a stomach divided into four compartments
salivary amylase
enzyme found in saliva, which converts carbohydrates to maltose
small intestine
organ where digestion of protein, fats, and carbohydrates is completed
sphincter
band of muscle that controls movement of materials throughout the digestive tract
stomach
saclike organ containing acidic digestive juices
villi
folds on the inner surface of the small intestine whose role is to increase absorption area
46.08: Variations in Vertebrate Digestive Systems
Skills to Develop
• Explain why an animal’s diet should be balanced and meet the needs of the body
• Define the primary components of food
• Describe the essential nutrients required for cellular function that cannot be synthesized by the animal body
• Explain how energy is produced through diet and digestion
• Describe how excess carbohydrates and energy are stored in the body
Given the diversity of animal life on our planet, it is not surprising that the animal diet would also vary substantially. The animal diet is the source of materials needed for building DNA and other complex molecules needed for growth, maintenance, and reproduction; collectively these processes are called biosynthesis. The diet is also the source of materials for ATP production in the cells. The diet must be balanced to provide the minerals and vitamins that are required for cellular function.
Food Requirements
What are the fundamental requirements of the animal diet? The animal diet should be well balanced and provide nutrients required for bodily function and the minerals and vitamins required for maintaining structure and regulation necessary for good health and reproductive capability. These requirements for a human are illustrated graphically in Figure \(1\).
Everyday Connection: Let’s Move! Campaign
Obesity is a growing epidemic and the rate of obesity among children is rapidly rising in the United States. To combat childhood obesity and ensure that children get a healthy start in life, first lady Michelle Obama has launched the Let’s Move! campaign. The goal of this campaign is to educate parents and caregivers on providing healthy nutrition and encouraging active lifestyles to future generations. This program aims to involve the entire community, including parents, teachers, and healthcare providers to ensure that children have access to healthy foods—more fruits, vegetables, and whole grains—and consume fewer calories from processed foods. Another goal is to ensure that children get physical activity. With the increase in television viewing and stationary pursuits such as video games, sedentary lifestyles have become the norm. Learn more at www.letsmove.gov.
Organic Precursors
The organic molecules required for building cellular material and tissues must come from food. Carbohydrates or sugars are the primary source of organic carbons in the animal body. During digestion, digestible carbohydrates are ultimately broken down into glucose and used to provide energy through metabolic pathways. Complex carbohydrates, including polysaccharides, can be broken down into glucose through biochemical modification; however, humans do not produce the enzyme cellulase and lack the ability to derive glucose from the polysaccharide cellulose. In humans, these molecules provide the fiber required for moving waste through the large intestine and a healthy colon. The intestinal flora in the human gut are able to extract some nutrition from these plant fibers. The excess sugars in the body are converted into glycogen and stored in the liver and muscles for later use. Glycogen stores are used to fuel prolonged exertions, such as long-distance running, and to provide energy during food shortage. Excess glycogen can be converted to fats, which are stored in the lower layer of the skin of mammals for insulation and energy storage. Excess digestible carbohydrates are stored by mammals in order to survive famine and aid in mobility.
Another important requirement is that of nitrogen. Protein catabolism provides a source of organic nitrogen. Amino acids are the building blocks of proteins and protein breakdown provides amino acids that are used for cellular function. The carbon and nitrogen derived from these become the building block for nucleotides, nucleic acids, proteins, cells, and tissues. Excess nitrogen must be excreted as it is toxic. Fats add flavor to food and promote a sense of satiety or fullness. Fatty foods are also significant sources of energy because one gram of fat contains nine calories. Fats are required in the diet to aid the absorption of fat-soluble vitamins and the production of fat-soluble hormones.
Essential Nutrients
While the animal body can synthesize many of the molecules required for function from the organic precursors, there are some nutrients that need to be consumed from food. These nutrients are termed essential nutrients, meaning they must be eaten, and the body cannot produce them.
The omega-3 alpha-linolenic acid and the omega-6 linoleic acid are essential fatty acids needed to make some membrane phospholipids. Vitamins are another class of essential organic molecules that are required in small quantities for many enzymes to function and, for this reason, are considered to be co-enzymes. Absence or low levels of vitamins can have a dramatic effect on health, as outlined in the tables below. Both fat-soluble and water-soluble vitamins must be obtained from food. Minerals, listed in the table below, are inorganic essential nutrients that must be obtained from food. Among their many functions, minerals help in structure and regulation and are considered co-factors. Certain amino acids also must be procured from food and cannot be synthesized by the body. These amino acids are the “essential” amino acids. The human body can synthesize only 11 of the 20 required amino acids; the rest must be obtained from food. The essential amino acids are listed in the table below.
Table \(1\): Water-soluble Essential Vitamins
Vitamin Function Deficiencies Can Lead To Sources
Vitamin B1 (Thiamine) Needed by the body to process lipids, proteins, and carbohydrates Coenzyme removes CO2 from organic compounds Muscle weakness, Beriberi: reduced heart function, CNS problems Milk, meat, dried beans, whole grains
Vitamin B2 (Riboflavin) Takes an active role in metabolism, aiding in the conversion of food to energy (FAD and FMN) Cracks or sores on the outer surface of the lips (cheliosis); inflammation and redness of the tongue; moist, scaly skin inflammation (seborrheic dermatitis) Meat, eggs, enriched grains, vegetables
Vitamin B3 (Niacin) Used by the body to release energy from carbohydrates and to process alcohol; required for the synthesis of sex hormones; component of coenzyme NAD+ and NADP+ Pellagra, which can result in dermatitis, diarrhea, dementia, and death Meat, eggs, grains, nuts, potatoes
Vitamin B5 (Pantothenic acid) Assists in producing energy from foods (lipids, in particular); component of coenzyme A Fatigue, poor coordination, retarded growth, numbness, tingling of hands and feet Meat, whole grains, milk, fruits, vegetables
Vitamin B6 (Pyridoxine) The principal vitamin for processing amino acids and lipids; also helps convert nutrients into energy Irritability, depression, confusion, mouth sores or ulcers, anemia, muscular twitching Meat, dairy products, whole grains, orange juice
Vitamin B7 (Biotin) Used in energy and amino acid metabolism, fat synthesis, and fat breakdown; helps the body use blood sugar Hair loss, dermatitis, depression, numbness and tingling in the extremities; neuromuscular disorders Meat, eggs, legumes and other vegetables
Vitamin B9 (Folic acid) Assists the normal development of cells, especially during fetal development; helps metabolize nucleic and amino acids Deficiency during pregnancy is associated with birth defects, such as neural tube defects and anemia Leafy green vegetables, whole wheat, fruits, nuts, legumes
Vitamin B12 (Cobalamin) Maintains healthy nervous system and assists with blood cell formation; coenzyme in nucleic acid metabolism Anemia, neurological disorders, numbness, loss of balance Meat, eggs, animal products
Vitamin C (Ascorbic acid) Helps maintain connective tissue: bone, cartilage, and dentin; boosts the immune system Scurvy, which results in bleeding, hair and tooth loss; joint pain and swelling; delayed wound healing Citrus fruits, broccoli, tomatoes, red sweet bell peppers
Table \(2\): Fat-soluble Essential Vitamins
Vitamin Function Deficiencies Can Lead To Sources
Vitamin A (Retinol) Critical to the development of bones, teeth, and skin; helps maintain eyesight, enhances the immune system, fetal development, gene expression Night-blindness, skin disorders, impaired immunity Dark green leafy vegetables, yellow-orange vegetables fruits, milk, butter
Vitamin D Critical for calcium absorption for bone development and strength; maintains a stable nervous system; maintains a normal and strong heartbeat; helps in blood clotting Rickets, osteomalacia, immunity Cod liver oil, milk, egg yolk
Vitamin E (Tocopherol) Lessens oxidative damage of cells,and prevents lung damage from pollutants; vital to the immune system Deficiency is rare; anemia, nervous system degeneration Wheat germ oil, unrefined vegetable oils, nuts, seeds, grains
Vitamin K (Phylloquinone) Essential to blood clotting Bleeding and easy bruising Leafy green vegetables, tea
Table \(3\): Minerals and Their Function in the Human Body
Mineral Function Deficiencies Can Lead To Sources
*Calcium Needed for muscle and neuron function; heart health; builds bone and supports synthesis and function of blood cells; nerve function Osteoporosis, rickets, muscle spasms, impaired growth Milk, yogurt, fish, green leafy vegetables, legumes
*Chlorine Needed for production of hydrochloric acid (HCl) in the stomach and nerve function; osmotic balance Muscle cramps, mood disturbances, reduced appetite Table salt
Copper (trace amounts) Required component of many redox enzymes, including cytochrome c oxidase; cofactor for hemoglobin synthesis Copper deficiency is rare Liver, oysters, cocoa, chocolate, sesame, nuts
Iodine Required for the synthesis of thyroid hormones Goiter Seafood, iodized salt, dairy products
Iron Required for many proteins and enzymes, notably hemoglobin, to prevent anemia Anemia, which causes poor concentration, fatigue, and poor immune function Red meat, leafy green vegetables, fish (tuna, salmon), eggs, dried fruits, beans, whole grains
*Magnesium Required co-factor for ATP formation; bone formation; normal membrane functions; muscle function Mood disturbances, muscle spasms Whole grains, leafy green vegetables
Manganese (trace amounts) A cofactor in enzyme functions; trace amounts are required Manganese deficiency is rare Common in most foods
Molybdenum (trace amounts) Acts as a cofactor for three essential enzymes in humans: sulfite oxidase, xanthine oxidase, and aldehyde oxidase Molybdenum deficiency is rare
*Phosphorus A component of bones and teeth; helps regulate acid-base balance; nucleotide synthesis Weakness, bone abnormalities, calcium loss Milk, hard cheese, whole grains, meats
*Potassium Vital for muscles, heart, and nerve function Cardiac rhythm disturbance, muscle weakness Legumes, potato skin, tomatoes, bananas
Selenium (trace amounts) A cofactor essential to activity of antioxidant enzymes like glutathione peroxidase; trace amounts are required Selenium deficiency is rare Common in most foods
*Sodium Systemic electrolyte required for many functions; acid-base balance; water balance; nerve function Muscle cramps, fatigue, reduced appetite Table salt
Zinc (trace amounts) Required for several enzymes such as carboxypeptidase, liver alcohol dehydrogenase, and carbonic anhydrase Anemia, poor wound healing, can lead to short stature Common in most foods
*Greater than 200mg/day required
Table \(4\): Essential Amino Acids
Amino acids that must be consumed Amino acids anabolized by the body
isoleucine alanine
leucine selenocysteine
lysine aspartate
methionine cysteine
phenylalanine glutamate
tryptophan glycine
valine proline
histidine* serine
threonine tyrosine
arginine* asparagine
*The human body can synthesize histidine and arginine, but not in the quantities required, especially for growing children.
Food Energy and ATP
Animals need food to obtain energy and maintain homeostasis. Homeostasis is the ability of a system to maintain a stable internal environment even in the face of external changes to the environment. For example, the normal body temperature of humans is 37°C (98.6°F). Humans maintain this temperature even when the external temperature is hot or cold. It takes energy to maintain this body temperature, and animals obtain this energy from food.
The primary source of energy for animals is carbohydrates, mainly glucose. Glucose is called the body’s fuel. The digestible carbohydrates in an animal’s diet are converted to glucose molecules through a series of catabolic chemical reactions.
Adenosine triphosphate, or ATP, is the primary energy currency in cells; ATP stores energy in phosphate ester bonds. ATP releases energy when the phosphodiester bonds are broken and ATP is converted to ADP and a phosphate group. ATP is produced by the oxidative reactions in the cytoplasm and mitochondrion of the cell, where carbohydrates, proteins, and fats undergo a series of metabolic reactions collectively called cellular respiration. For example, glycolysis is a series of reactions in which glucose is converted to pyruvic acid and some of its chemical potential energy is transferred to NADH and ATP.
ATP is required for all cellular functions. It is used to build the organic molecules that are required for cells and tissues; it provides energy for muscle contraction and for the transmission of electrical signals in the nervous system. When the amount of ATP is available in excess of the body’s requirements, the liver uses the excess ATP and excess glucose to produce molecules called glycogen. Glycogen is a polymeric form of glucose and is stored in the liver and skeletal muscle cells. When blood sugar drops, the liver releases glucose from stores of glycogen. Skeletal muscle converts glycogen to glucose during intense exercise. The process of converting glucose and excess ATP to glycogen and the storage of excess energy is an evolutionarily important step in helping animals deal with mobility, food shortages, and famine.
Everyday Connection: Obesity
Obesity is a major health concern in the United States, and there is a growing focus on reducing obesity and the diseases it may lead to, such as type-2 diabetes, cancers of the colon and breast, and cardiovascular disease. How does the food consumed contribute to obesity?
Fatty foods are calorie-dense, meaning that they have more calories per unit mass than carbohydrates or proteins. One gram of carbohydrates has four calories, one gram of protein has four calories, and one gram of fat has nine calories. Animals tend to seek lipid-rich food for their higher energy content.
The signals of hunger (“time to eat”) and satiety (“time to stop eating”) are controlled in the hypothalamus region of the brain. Foods that are rich in fatty acids tend to promote satiety more than foods that are rich only in carbohydrates.
Excess carbohydrate and ATP are used by the liver to synthesize glycogen. The pyruvate produced during glycolysis is used to synthesize fatty acids. When there is more glucose in the body than required, the resulting excess pyruvate is converted into molecules that eventually result in the synthesis of fatty acids within the body. These fatty acids are stored in adipose cells—the fat cells in the mammalian body whose primary role is to store fat for later use.
It is important to note that some animals benefit from obesity. Polar bears and seals need body fat for insulation and to keep them from losing body heat during Arctic winters. When food is scarce, stored body fat provides energy for maintaining homeostasis. Fats prevent famine in mammals, allowing them to access energy when food is not available on a daily basis; fats are stored when a large kill is made or lots of food is available.
Summary
Animal diet should be balanced and meet the needs of the body. Carbohydrates, proteins, and fats are the primary components of food. Some essential nutrients are required for cellular function but cannot be produced by the animal body. These include vitamins, minerals, some fatty acids, and some amino acids. Food intake in more than necessary amounts is stored as glycogen in the liver and muscle cells, and in fat cells. Excess adipose storage can lead to obesity and serious health problems. ATP is the energy currency of the cell and is obtained from the metabolic pathways. Excess carbohydrates and energy are stored as glycogen in the body.
Glossary
essential nutrient
nutrient that cannot be synthesized by the body; it must be obtained from food
mineral
inorganic, elemental molecule that carries out important roles in the body
vitamin
organic substance necessary in small amounts to sustain life | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/46%3A_The_Digestive_System/46.08%3A_Variations_in_Vertebrate_Digestive_Systems/46.8.01%3A_Nutrition_and_Energy_Production.txt |
• 47.1: Gas Exchange Across Respiratory Surfaces
• 47.2: Gills, Cutaneous Respirate, and Tracheal Systems
Modern fishes include an estimated 31,000 species. Fishes were the earliest vertebrates, with jawless species being the earliest and jawed species evolving later. They are active feeders, rather than sessile, suspension feeders. Jawless fishes—the hagfishes and lampreys—have a distinct cranium and complex sense organs including eyes, distinguishing them from the invertebrate chordates.
• 47.3: Lungs
The structure of the lung maximizes its surface area to increase gas diffusion. Because of the enormous number of alveoli (approximately 300 million in each human lung), the surface area of the lung is very large (75 square meters). Having such a large surface area increases the amount of gas that can diffuse into and out of the lungs.
• 47.4: Structures, Mechanisms, and Control of Ventilation in Mammals
• 47.5: Transport of Gases in Body Fluid
Once the oxygen diffuses across the alveoli, it enters the bloodstream and is transported to the tissues where it is unloaded, and carbon dioxide diffuses out of the blood and into the alveoli to be expelled from the body. Although gas exchange is a continuous process, the oxygen and carbon dioxide are transported by different mechanisms.
47: The Respiratory System
Skills to Develop
• Describe the difference between jawless and jawed fishes
• Discuss the distinguishing features of sharks and rays compared to other modern fishes
Modern fishes include an estimated 31,000 species. Fishes were the earliest vertebrates, with jawless species being the earliest and jawed species evolving later. They are active feeders, rather than sessile, suspension feeders. Jawless fishes—the hagfishes and lampreys—have a distinct cranium and complex sense organs including eyes, distinguishing them from the invertebrate chordates.
Jawless Fishes
Jawless fishes are craniates that represent an ancient vertebrate lineage that arose over one half-billion years ago. In the past, the hagfishes and lampreys were classified together as agnathans. Today, hagfishes and lampreys are recognized as separate clades, primarily because lampreys are true vertebrates, whereas hagfishes are not. A defining feature is the lack of paired lateral appendages (fins). Some of the earliest jawless fishes were the ostracoderms (which translates to “shell-skin”). Ostracoderms were vertebrate fishes encased in bony armor, unlike present-day jawless fishes, which lack bone in their scales.
Myxini: Hagfishes
The clade Myxini includes at least 20 species of hagfishes. Hagfishes are eel-like scavengers that live on the ocean floor and feed on dead invertebrates, other fishes, and marine mammals (Figure \(1\)). Hagfishes are entirely marine and are found in oceans around the world, except for the polar regions. A unique feature of these animals is the slime glands beneath the skin that release mucus through surface pores. This mucus allows the hagfish to escape from the grip of predators. Hagfish can also twist their bodies in a knot to feed and sometimes eat carcasses from the inside out.
The skeleton of a hagfish is composed of cartilage, which includes a cartilaginous notochord that runs the length of the body. This notochord provides support to the hagfish’s body. Hagfishes do not replace the notochord with a vertebral column during development, as do true vertebrates.
Petromyzontidae: Lampreys
The clade Petromyzontidae includes approximately 35–40 or more species of lampreys. Lampreys are similar to hagfishes in size and shape; however, lampreys possess some vertebral elements. Lampreys lack paired appendages and bone, as do the hagfishes. As adults, lampreys are characterized by a toothed, funnel-like sucking mouth. Many species have a parasitic stage of their life cycle during which they are ectoparasites of fishes (Figure \(2\)).
Lampreys live primarily in coastal and fresh waters, and have a worldwide distribution, except for in the tropics and polar regions. Some species are marine, but all species spawn in fresh water. Eggs are fertilized externally, and the larvae distinctly differ from the adult form, spending 3 to 15 years as suspension feeders. Once they attain sexual maturity, the adults reproduce and die within days.
Lampreys possess a notochord as adults; however, this notochord is surrounded by a cartilaginous structure called an arcualia, which may resemble an evolutionarily early form of the vertebral column.
Gnathostomes: Jawed Fishes
Gnathostomes or “jaw-mouths” are vertebrates that possess jaws. One of the most significant developments in early vertebrate evolution was the development of the jaw, which is a hinged structure attached to the cranium that allows an animal to grasp and tear its food. The evolution of jaws allowed early gnathostomes to exploit food resources that were unavailable to jawless fishes.
Early gnathostomes also possessed two sets of paired fins, allowing the fishes to maneuver accurately. Pectoral fins are typically located on the anterior body, and pelvic fins on the posterior. Evolution of the jaw and paired fins permitted gnathostomes to expand from the sedentary suspension feeding of jawless fishes to become mobile predators. The ability of gnathostomes to exploit new nutrient sources likely is one reason that they replaced most jawless fishes during the Devonian period. Two early groups of gnathostomes were the acanthodians and placoderms (Figure \(3\)), which arose in the late Silurian period and are now extinct. Most modern fishes are gnathostomes that belong to the clades Chondrichthyes and Osteichthyes.
Chondrichthyes: Cartilaginous Fishes
The clade Chondrichthyes is diverse, consisting of sharks (Figure \(4\)), rays, and skates, together with sawfishes and a few dozen species of fishes called chimaeras, or “ghost” sharks.” Chondrichthyes are jawed fishes that possess paired fins and a skeleton made of cartilage. This clade arose approximately 370 million years ago in the early or middle Devonian. They are thought to be descended from the placoderms, which had skeletons made of bone; thus, the cartilaginous skeleton of Chondrichthyes is a later development. Parts of shark skeleton are strengthened by granules of calcium carbonate, but this is not the same as bone.
Most cartilaginous fishes live in marine habitats, with a few species living in fresh water for a part or all of their lives. Most sharks are carnivores that feed on live prey, either swallowing it whole or using their jaws and teeth to tear it into smaller pieces. Shark teeth likely evolved from the jagged scales that cover their skin, called placoid scales. Some species of sharks and rays are suspension feeders that feed on plankton.
Sharks have well-developed sense organs that aid them in locating prey, including a keen sense of smell and electroreception, with the latter perhaps the most sensitive of any animal. Organs called ampullae of Lorenzini allow sharks to detect the electromagnetic fields that are produced by all living things, including their prey. Electroreception has only been observed in aquatic or amphibious animals. Sharks, together with most fishes and aquatic and larval amphibians, also have a sense organ called the lateral line, which is used to detect movement and vibration in the surrounding water, and is often considered homologous to “hearing” in terrestrial vertebrates. The lateral line is visible as a darker stripe that runs along the length of a fish’s body.
Sharks reproduce sexually, and eggs are fertilized internally. Most species are ovoviviparous: The fertilized egg is retained in the oviduct of the mother’s body and the embryo is nourished by the egg yolk. The eggs hatch in the uterus, and young are born alive and fully functional. Some species of sharks are oviparous: They lay eggs that hatch outside of the mother’s body. Embryos are protected by a shark egg case or “mermaid’s purse” (Figure \(5\)) that has the consistency of leather. The shark egg case has tentacles that snag in seaweed and give the newborn shark cover. A few species of sharks are viviparous: The young develop within the mother’s body and she gives live birth.
Rays and skates comprise more than 500 species and are closely related to sharks. They can be distinguished from sharks by their flattened bodies, pectoral fins that are enlarged and fused to the head, and gill slits on their ventral surface (Figure \(6\)). Like sharks, rays and skates have a cartilaginous skeleton. Most species are marine and live on the sea floor, with nearly a worldwide distribution.
Osteichthyes: Bony Fishes
Members of the clade Osteichthyes, also called bony fishes, are characterized by a bony skeleton. The vast majority of present-day fishes belong to this group, which consists of approximately 30,000 species, making it the largest class of vertebrates in existence today.
Nearly all bony fishes have an ossified skeleton with specialized bone cells (osteocytes) that produce and maintain a calcium phosphate matrix. This characteristic has only reversed in a few groups of Osteichthyes, such as sturgeons and paddlefish, which have primarily cartilaginous skeletons. The skin of bony fishes is often covered by overlapping scales, and glands in the skin secrete mucus that reduces drag when swimming and aids the fish in osmoregulation. Like sharks, bony fishes have a lateral line system that detects vibrations in water.
All bony fishes use gills to breathe. Water is drawn over gills that are located in chambers covered and ventilated by a protective, muscular flap called the operculum. Many bony fishes also have a swim bladder, a gas-filled organ that helps to control the buoyancy of the fish. Bony fishes are further divided into two extant clades: Actinopterygii (ray-finned fishes) and Sarcopterygii (lobe-finned fishes).
Actinopterygii, the ray-finned fishes, include many familiar fishes—tuna, bass, trout, and salmon (Figure \(7\)), among others. Ray-finned fishes are named for their fins that are webs of skin supported by bony spines called rays. In contrast, the fins of Sarcopterygii are fleshy and lobed, supported by bone (Figure \(7\)). Living members of this clade include the less-familiar lungfishes and coelacanths.
Summary
The earliest vertebrates that diverged from the invertebrate chordates were the jawless fishes. Fishes with jaws (gnathostomes) evolved later. Jaws allowed early gnathostomes to exploit new food sources. Agnathans include the hagfishes and lampreys. Hagfishes are eel-like scavengers that feed on dead invertebrates and other fishes. Lampreys are characterized by a toothed, funnel-like sucking mouth, and most species are parasitic on other fishes. Gnathostomes include the cartilaginous fishes and the bony fishes, as well as all other tetrapods. Cartilaginous fishes include sharks, rays, skates, and ghost sharks. Most cartilaginous fishes live in marine habitats, with a few species living in fresh water for part or all of their lives. The vast majority of present-day fishes belong to the clade Osteichthyes, which consists of approximately 30,000 species. Bony fishes can be divided into two clades: Actinopterygii (ray-finned fishes, virtually all extant species) and Sarcopterygii (lobe-finned fishes, comprising fewer than 10 extant species but which are the ancestors of tetrapods).
Glossary
Actinopterygii
ray-finned fishes
ampulla of Lorenzini
sensory organ that allows sharks to detect electromagnetic fields produced by living things
Chondrichthyes
jawed fish with paired fins and a skeleton made of cartilage
gnathostome
jawed fish
hagfish
eel-like jawless fish that live on the ocean floor and are scavengers
lamprey
jawless fish characterized by a toothed, funnel-like, sucking mouth
lateral line
sense organ that runs the length of a fish’s body; used to detect vibration in the water
Myxini
hagfishes
Osteichthyes
bony fish
ostracoderm
one of the earliest jawless fish covered in bone
Petromyzontidae
clade of lampreys
Sarcopterygii
lobe-finned fish
swim bladder
in fishes, a gas filled organ that helps to control the buoyancy of the fish
47.02: Gills Cutaneous Respirate and Tracheal Systems
Skills to Develop
• Describe the important difference between the life cycle of amphibians and the life cycles of other vertebrates
• Distinguish between the characteristics of Urodela, Anura, and Apoda
• Describe the evolutionary history of amphibians
Amphibians are vertebrate tetrapods. Amphibia includes frogs, salamanders, and caecilians. The term amphibian loosely translates from the Greek as “dual life,” which is a reference to the metamorphosis that many frogs and salamanders undergo and their mixture of aquatic and terrestrial environments in their life cycle. Amphibians evolved during the Devonian period and were the earliest terrestrial tetrapods.
Characteristics of Amphibians
As tetrapods, most amphibians are characterized by four well-developed limbs. Some species of salamanders and all caecilians are functionally limbless; their limbs are vestigial. An important characteristic of extant amphibians is a moist, permeable skin that is achieved via mucus glands that keep the skin moist; thus, exchange of oxygen and carbon dioxide with the environment can take place through it (cutaneous respiration). Additional characteristics of amphibians include pedicellate teeth—teeth in which the root and crown are calcified, separated by a zone of noncalcified tissue—and a papilla amphibiorum and papilla basilaris, structures of the inner ear that are sensitive to frequencies below and above 10,00 hertz, respectively. Amphibians also have an auricular operculum, which is an extra bone in the ear that transmits sounds to the inner ear. All extant adult amphibians are carnivorous, and some terrestrial amphibians have a sticky tongue that is used to capture prey.
Evolution of Amphibians
The fossil record provides evidence of the first tetrapods: now-extinct amphibian species dating to nearly 400 million years ago. Evolution of tetrapods from fishes represented a significant change in body plan from one suited to organisms that respired and swam in water, to organisms that breathed air and moved onto land; these changes occurred over a span of 50 million years during the Devonian period. One of the earliest known tetrapods is from the genus Acanthostega. Acanthostega was aquatic; fossils show that it had gills similar to fishes. However, it also had four limbs, with the skeletal structure of limbs found in present-day tetrapods, including amphibians. Therefore, it is thought that Acanthostega lived in shallow waters and was an intermediate form between lobe-finned fishes and early, fully terrestrial tetrapods. What preceded Acanthostega?
In 2006, researchers published news of their discovery of a fossil of a “tetrapod-like fish,” Tiktaalik roseae, which seems to be an intermediate form between fishes having fins and tetrapods having limbs (Figure \(1\)). Tiktaalik likely lived in a shallow water environment about 375 million years ago.1
The early tetrapods that moved onto land had access to new nutrient sources and relatively few predators. This led to the widespread distribution of tetrapods during the early Carboniferous period, a period sometimes called the “age of the amphibians.”
Modern Amphibians
Amphibia comprises an estimated 6,770 extant species that inhabit tropical and temperate regions around the world. Amphibians can be divided into three clades: Urodela (“tailed-ones”), the salamanders; Anura (“tail-less ones”), the frogs; and Apoda (“legless ones”), the caecilians.
Urodela: Salamanders
Salamanders are amphibians that belong to the order Urodela. Living salamanders (Figure \(1\)) include approximately 620 species, some of which are aquatic, other terrestrial, and some that live on land only as adults. Adult salamanders usually have a generalized tetrapod body plan with four limbs and a tail. They move by bending their bodies from side to side, called lateral undulation, in a fish-like manner while “walking” their arms and legs fore and aft. It is thought that their gait is similar to that used by early tetrapods. Respiration differs among different species. The majority of salamanders are lungless, and respiration occurs through the skin or through external gills. Some terrestrial salamanders have primitive lungs; a few species have both gills and lungs.
Unlike frogs, virtually all salamanders rely on internal fertilization of the eggs. The only male amphibians that possess copulatory structures are the caecilians, so fertilization among salamanders typically involves an elaborate and often prolonged courtship. Such a courtship allows the successful transfer of sperm from male to female via a spermatophore. Development in many of the most highly evolved salamanders, which are fully terrestrial, occurs during a prolonged egg stage, with the eggs guarded by the mother. During this time, the gilled larval stage is found only within the egg capsule, with the gills being resorbed, and metamorphosis being completed, before hatching. Hatchlings thus resemble tiny adults.
Link to Learning
View River Monsters: Fish With Arms and Hands? to see a video about an unusually large salamander species.
Anura: Frogs
Frogs are amphibians that belong to the order Anura (Figure \(3\)). Anurans are among the most diverse groups of vertebrates, with approximately 5,965 species that occur on all of the continents except Antarctica. Anurans have a body plan that is more specialized for movement. Adult frogs use their hind limbs to jump on land. Frogs have a number of modifications that allow them to avoid predators, including skin that acts as camouflage. Many species of frogs and salamanders also release defensive chemicals from glands in the skin that are poisonous to predators.
Frog eggs are fertilized externally, and like other amphibians, frogs generally lay their eggs in moist environments. A moist environment is required as eggs lack a shell and thus dehydrate quickly in dry environments. Frogs demonstrate a great diversity of parental behaviors, with some species laying many eggs and exhibiting little parental care, to species that carry eggs and tadpoles on their hind legs or backs. The life cycle of frogs, as other amphibians, consists of two distinct stages: the larval stage followed by metamorphosis to an adult stage. The larval stage of a frog, the tadpole, is often a filter-feeding herbivore. Tadpoles usually have gills, a lateral line system, long-finned tails, and lack limbs. At the end of the tadpole stage, frogs undergo metamorphosis into the adult form (Figure \(4\)). During this stage, the gills, tail, and lateral line system disappear, and four limbs develop. The jaws become larger and are suited for carnivorous feeding, and the digestive system transforms into the typical short gut of a predator. An eardrum and air-breathing lungs also develop. These changes during metamorphosis allow the larvae to move onto land in the adult stage.
Apoda: Caecilians
An estimated 185 species comprise caecilians, a group of amphibians that belong to the order Apoda. Although they are vertebrates, a complete lack of limbs leads to their resemblance to earthworms in appearance. They are adapted for a soil-burrowing or aquatic lifestyle, and they are nearly blind. These animals are found in the tropics of South America, Africa, and Southern Asia. They have vestigial limbs, evidence that they evolved from a legged ancestor.
Evolution Connection: The Paleozoic Era and the Evolution of Vertebrates
The climate and geography of Earth was vastly different during the Paleozoic Era, when vertebrates arose, as compared to today. The Paleozoic spanned from approximately 542 to 251 million years ago. The landmasses on Earth were very different from those of today. Laurentia and Gondwana were continents located near the equator that subsumed much of the current day landmasses in a different configuration (Figure \(5\)). At this time, sea levels were very high, probably at a level that hasn’t been reached since. As the Paleozoic progressed, glaciations created a cool global climate, but conditions warmed near the end of the first half of the Paleozoic. During the latter half of the Paleozoic, the landmasses began moving together, with the initial formation of a large northern block called Laurasia. This contained parts of what is now North America, along with Greenland, parts of Europe, and Siberia. Eventually, a single supercontinent, called Pangaea, was formed, starting in the latter third of the Paleozoic. Glaciations then began to affect Pangaea’s climate, affecting the distribution of vertebrate life.
During the early Paleozoic, the amount of carbon dioxide in the atmosphere was much greater than it is today. This may have begun to change later, as land plants became more common. As the roots of land plants began to infiltrate rock and soil began to form, carbon dioxide was drawn out of the atmosphere and became trapped in the rock. This reduced the levels of carbon dioxide and increased the levels of oxygen in the atmosphere, so that by the end of the Paleozoic, atmospheric conditions were similar to those of today.
As plants became more common through the latter half of the Paleozoic, microclimates began to emerge and ecosystems began to change. As plants and ecosystems continued to grow and become more complex, vertebrates moved from the water to land. The presence of shoreline vegetation may have contributed to the movement of vertebrates onto land. One hypothesis suggests that the fins of aquatic vertebrates were used to maneuver through this vegetation, providing a precursor to the movement of fins on land and the development of limbs. The late Paleozoic was a time of diversification of vertebrates, as amniotes emerged and became two different lines that gave rise, on one hand, to mammals, and, on the other hand, to reptiles and birds. Many marine vertebrates became extinct near the end of the Devonian period, which ended about 360 million years ago, and both marine and terrestrial vertebrates were decimated by a mass extinction in the early Permian period about 250 million years ago.
Link to Learning
View Earth’s Paleogeography: Continental Movements Through Time to see changes in Earth as life evolved.
Summary
As tetrapods, most amphibians are characterized by four well-developed limbs, although some species of salamanders and all caecilians are limbless. The most important characteristic of extant amphibians is a moist, permeable skin used for cutaneous respiration. The fossil record provides evidence of amphibian species, now extinct, that arose over 400 million years ago as the first tetrapods. Amphibia can be divided into three clades: salamanders (Urodela), frogs (Anura), and caecilians (Apoda). The life cycle of frogs, like the majority of amphibians, consists of two distinct stages: the larval stage and metamorphosis to an adult stage. Some species in all orders bypass a free-living larval stage.
Footnotes
1. 1 Daeschler, E. B., Shubin, N. H., and Jenkins, F. J. “A Devonian tetrapod-like fish and the evolution of the tetrapod body plan,” Nature 440 (2006): 757–763, doi:10.1038/nature04639, http://www.nature.com/nature/journal/v440/n7085/abs/nature04639.html.
Glossary
Acanthostega
one of the earliest known tetrapods
Amphibia
frogs, salamanders, and caecilians
Anura
frogs
Apoda
caecilians
caecilian
legless amphibian that belongs to the clade Apoda
cutaneous respiration
gas exchange through the skin
frog
tail-less amphibian that belongs to the clade Anura
salamander
tailed amphibian that belongs to the clade Urodela
tadpole
larval stage of a frog
Urodela
salamanders | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/47%3A_The_Respiratory_System/47.02%3A_Gills_Cutaneous_Respirate_and_Tracheal_Systems/47.2.01%3A_Amphibians.txt |
Skills to Develop
• Describe the passage of air from the outside environment to the lungs
• Explain how the lungs are protected from particulate matter
The primary function of the respiratory system is to deliver oxygen to the cells of the body’s tissues and remove carbon dioxide, a cell waste product. The main structures of the human respiratory system are the nasal cavity, the trachea, and lungs.
All aerobic organisms require oxygen to carry out their metabolic functions. Along the evolutionary tree, different organisms have devised different means of obtaining oxygen from the surrounding atmosphere. The environment in which the animal lives greatly determines how an animal respires. The complexity of the respiratory system is correlated with the size of the organism. As animal size increases, diffusion distances increase and the ratio of surface area to volume drops. In unicellular organisms, diffusion across the cell membrane is sufficient for supplying oxygen to the cell (Figure \(1\)). Diffusion is a slow, passive transport process. In order for diffusion to be a feasible means of providing oxygen to the cell, the rate of oxygen uptake must match the rate of diffusion across the membrane. In other words, if the cell were very large or thick, diffusion would not be able to provide oxygen quickly enough to the inside of the cell. Therefore, dependence on diffusion as a means of obtaining oxygen and removing carbon dioxide remains feasible only for small organisms or those with highly-flattened bodies, such as many flatworms (Platyhelminthes). Larger organisms had to evolve specialized respiratory tissues, such as gills, lungs, and respiratory passages accompanied by complex circulatory systems, to transport oxygen throughout their entire body.
Direct Diffusion
For small multicellular organisms, diffusion across the outer membrane is sufficient to meet their oxygen needs. Gas exchange by direct diffusion across surface membranes is efficient for organisms less than 1 mm in diameter. In simple organisms, such as cnidarians and flatworms, every cell in the body is close to the external environment. Their cells are kept moist and gases diffuse quickly via direct diffusion. Flatworms are small, literally flat worms, which ‘breathe’ through diffusion across the outer membrane (Figure \(2\)). The flat shape of these organisms increases the surface area for diffusion, ensuring that each cell within the body is close to the outer membrane surface and has access to oxygen. If the flatworm had a cylindrical body, then the cells in the center would not be able to get oxygen.
Skin and Gills
Earthworms and amphibians use their skin (integument) as a respiratory organ. A dense network of capillaries lies just below the skin and facilitates gas exchange between the external environment and the circulatory system. The respiratory surface must be kept moist in order for the gases to dissolve and diffuse across cell membranes.
Organisms that live in water need to obtain oxygen from the water. Oxygen dissolves in water but at a lower concentration than in the atmosphere. The atmosphere has roughly 21 percent oxygen. In water, the oxygen concentration is much smaller than that. Fish and many other aquatic organisms have evolved gills to take up the dissolved oxygen from water (Figure \(3\)). Gills are thin tissue filaments that are highly branched and folded. When water passes over the gills, the dissolved oxygen in water rapidly diffuses across the gills into the bloodstream. The circulatory system can then carry the oxygenated blood to the other parts of the body. In animals that contain coelomic fluid instead of blood, oxygen diffuses across the gill surfaces into the coelomic fluid. Gills are found in mollusks, annelids, and crustaceans.
The folded surfaces of the gills provide a large surface area to ensure that the fish gets sufficient oxygen. Diffusion is a process in which material travels from regions of high concentration to low concentration until equilibrium is reached. In this case, blood with a low concentration of oxygen molecules circulates through the gills. The concentration of oxygen molecules in water is higher than the concentration of oxygen molecules in gills. As a result, oxygen molecules diffuse from water (high concentration) to blood (low concentration), as shown in Figure \(4\). Similarly, carbon dioxide molecules in the blood diffuse from the blood (high concentration) to water (low concentration).
Tracheal Systems
Insect respiration is independent of its circulatory system; therefore, the blood does not play a direct role in oxygen transport. Insects have a highly specialized type of respiratory system called the tracheal system, which consists of a network of small tubes that carries oxygen to the entire body. The tracheal system is the most direct and efficient respiratory system in active animals. The tubes in the tracheal system are made of a polymeric material called chitin.
Insect bodies have openings, called spiracles, along the thorax and abdomen. These openings connect to the tubular network, allowing oxygen to pass into the body (Figure \(5\)) and regulating the diffusion of CO2 and water vapor. Air enters and leaves the tracheal system through the spiracles. Some insects can ventilate the tracheal system with body movements.
Mammalian Systems
In mammals, pulmonary ventilation occurs via inhalation (breathing). During inhalation, air enters the body through the nasal cavity located just inside the nose (Figure \(6\)). As air passes through the nasal cavity, the air is warmed to body temperature and humidified. The respiratory tract is coated with mucus to seal the tissues from direct contact with air. Mucus is high in water. As air crosses these surfaces of the mucous membranes, it picks up water. These processes help equilibrate the air to the body conditions, reducing any damage that cold, dry air can cause. Particulate matter that is floating in the air is removed in the nasal passages via mucus and cilia. The processes of warming, humidifying, and removing particles are important protective mechanisms that prevent damage to the trachea and lungs. Thus, inhalation serves several purposes in addition to bringing oxygen into the respiratory system.
Art Connection
Which of the following statements about the mammalian respiratory system is false?
1. When we breathe in, air travels from the pharynx to the trachea.
2. The bronchioles branch into bronchi.
3. Alveolar ducts connect to alveolar sacs.
4. Gas exchange between the lung and blood takes place in the alveolus.
From the nasal cavity, air passes through the pharynx (throat) and the larynx (voice box), as it makes its way to the trachea (Figure \(6\)). The main function of the trachea is to funnel the inhaled air to the lungs and the exhaled air back out of the body. The human trachea is a cylinder about 10 to 12 cm long and 2 cm in diameter that sits in front of the esophagus and extends from the larynx into the chest cavity where it divides into the two primary bronchi at the midthorax. It is made of incomplete rings of hyaline cartilage and smooth muscle (Figure \(7\)). The trachea is lined with mucus-producing goblet cells and ciliated epithelia. The cilia propel foreign particles trapped in the mucus toward the pharynx. The cartilage provides strength and support to the trachea to keep the passage open. The smooth muscle can contract, decreasing the trachea’s diameter, which causes expired air to rush upwards from the lungs at a great force. The forced exhalation helps expel mucus when we cough. Smooth muscle can contract or relax, depending on stimuli from the external environment or the body’s nervous system.
Lungs: Bronchi and Alveoli
The end of the trachea bifurcates (divides) to the right and left lungs. The lungs are not identical. The right lung is larger and contains three lobes, whereas the smaller left lung contains two lobes (Figure \(8\)). The muscular diaphragm, which facilitates breathing, is inferior to (below) the lungs and marks the end of the thoracic cavity.
In the lungs, air is diverted into smaller and smaller passages, or bronchi. Air enters the lungs through the two primary (main) bronchi (singular: bronchus). Each bronchus divides into secondary bronchi, then into tertiary bronchi, which in turn divide, creating smaller and smaller diameter bronchioles as they split and spread through the lung. Like the trachea, the bronchi are made of cartilage and smooth muscle. At the bronchioles, the cartilage is replaced with elastic fibers. Bronchi are innervated by nerves of both the parasympathetic and sympathetic nervous systems that control muscle contraction (parasympathetic) or relaxation (sympathetic) in the bronchi and bronchioles, depending on the nervous system’s cues. In humans, bronchioles with a diameter smaller than 0.5 mm are the respiratory bronchioles. They lack cartilage and therefore rely on inhaled air to support their shape. As the passageways decrease in diameter, the relative amount of smooth muscle increases.
The terminal bronchioles subdivide into microscopic branches called respiratory bronchioles. The respiratory bronchioles subdivide into several alveolar ducts. Numerous alveoli and alveolar sacs surround the alveolar ducts. The alveolar sacs resemble bunches of grapes tethered to the end of the bronchioles (Figure \(9\)). In the acinar region, the alveolar ducts are attached to the end of each bronchiole. At the end of each duct are approximately 100 alveolar sacs, each containing 20 to 30 alveoli that are 200 to 300 microns in diameter. Gas exchange occurs only in alveoli. Alveoli are made of thin-walled parenchymal cells, typically one-cell thick, that look like tiny bubbles within the sacs. Alveoli are in direct contact with capillaries (one-cell thick) of the circulatory system. Such intimate contact ensures that oxygen will diffuse from alveoli into the blood and be distributed to the cells of the body. In addition, the carbon dioxide that was produced by cells as a waste product will diffuse from the blood into alveoli to be exhaled. The anatomical arrangement of capillaries and alveoli emphasizes the structural and functional relationship of the respiratory and circulatory systems. Because there are so many alveoli (~300 million per lung) within each alveolar sac and so many sacs at the end of each alveolar duct, the lungs have a sponge-like consistency. This organization produces a very large surface area that is available for gas exchange. The surface area of alveoli in the lungs is approximately 75 m2. This large surface area, combined with the thin-walled nature of the alveolar parenchymal cells, allows gases to easily diffuse across the cells.
Link to Learning
Watch the following video to review the respiratory system.
Protective Mechanisms
The air that organisms breathe contains particulate matter such as dust, dirt, viral particles, and bacteria that can damage the lungs or trigger allergic immune responses. The respiratory system contains several protective mechanisms to avoid problems or tissue damage. In the nasal cavity, hairs and mucus trap small particles, viruses, bacteria, dust, and dirt to prevent their entry.
If particulates do make it beyond the nose, or enter through the mouth, the bronchi and bronchioles of the lungs also contain several protective devices. The lungs produce mucus—a sticky substance made of mucin, a complex glycoprotein, as well as salts and water—that traps particulates. The bronchi and bronchioles contain cilia, small hair-like projections that line the walls of the bronchi and bronchioles (Figure \(10\)). These cilia beat in unison and move mucus and particles out of the bronchi and bronchioles back up to the throat where it is swallowed and eliminated via the esophagus.
In humans, for example, tar and other substances in cigarette smoke destroy or paralyze the cilia, making the removal of particles more difficult. In addition, smoking causes the lungs to produce more mucus, which the damaged cilia are not able to move. This causes a persistent cough, as the lungs try to rid themselves of particulate matter, and makes smokers more susceptible to respiratory ailments.
Summary
Animal respiratory systems are designed to facilitate gas exchange. In mammals, air is warmed and humidified in the nasal cavity. Air then travels down the pharynx, through the trachea, and into the lungs. In the lungs, air passes through the branching bronchi, reaching the respiratory bronchioles, which house the first site of gas exchange. The respiratory bronchioles open into the alveolar ducts, alveolar sacs, and alveoli. Because there are so many alveoli and alveolar sacs in the lung, the surface area for gas exchange is very large. Several protective mechanisms are in place to prevent damage or infection. These include the hair and mucus in the nasal cavity that trap dust, dirt, and other particulate matter before they can enter the system. In the lungs, particles are trapped in a mucus layer and transported via cilia up to the esophageal opening at the top of the trachea to be swallowed.
Art Connections
Figure \(6\): Which of the following statements about the mammalian respiratory system is false?
1. When we breathe in, air travels from the pharynx to the trachea.
2. The bronchioles branch into bronchi.
3. Alveolar ducts connect to alveolar sacs.
4. Gas exchange between the lung and blood takes place in the alveolus.
Answer
B
Glossary
alveolar duct
duct that extends from the terminal bronchiole to the alveolar sac
alveolar sac
structure consisting of two or more alveoli that share a common opening
alveolus
(plural: alveoli) (also, air sac) terminal region of the lung where gas exchange occurs
bronchus
(plural: bronchi) smaller branch of cartilaginous tissue that stems off of the trachea; air is funneled through the bronchi to the region where gas exchange occurs in alveoli
bronchiole
airway that extends from the main tertiary bronchi to the alveolar sac
diaphragm
domed-shaped skeletal muscle located under lungs that separates the thoracic cavity from the abdominal cavity
larynx
voice box, a short passageway connecting the pharynx and the trachea
mucin
complex glycoprotein found in mucus
mucus
sticky protein-containing fluid secretion in the lung that traps particulate matter to be expelled from the body
nasal cavity
opening of the respiratory system to the outside environment
particulate matter
small particle such as dust, dirt, viral particles, and bacteria that are in the air
pharynx
throat; a tube that starts in the internal nares and runs partway down the neck, where it opens into the esophagus and the larynx
primary bronchus
(also, main bronchus) region of the airway within the lung that attaches to the trachea and bifurcates to each lung where it branches into secondary bronchi
respiratory bronchiole
terminal portion of the bronchiole tree that is attached to the terminal bronchioles and alveoli ducts, alveolar sacs, and alveoli
terminal bronchiole
region of bronchiole that attaches to the respiratory bronchioles
trachea
cartilaginous tube that transports air from the larynx to the primary bronchi | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/47%3A_The_Respiratory_System/47.02%3A_Gills_Cutaneous_Respirate_and_Tracheal_Systems/47.2.02%3A_Systems_of_Gas_Exchange.txt |
Skills to Develop
• Name and describe lung volumes and capacities
• Understand how gas pressure influences how gases move into and out of the body
The structure of the lung maximizes its surface area to increase gas diffusion. Because of the enormous number of alveoli (approximately 300 million in each human lung), the surface area of the lung is very large (75 m2). Having such a large surface area increases the amount of gas that can diffuse into and out of the lungs.
Basic Principles of Gas Exchange
Gas exchange during respiration occurs primarily through diffusion. Diffusion is a process in which transport is driven by a concentration gradient. Gas molecules move from a region of high concentration to a region of low concentration. Blood that is low in oxygen concentration and high in carbon dioxide concentration undergoes gas exchange with air in the lungs. The air in the lungs has a higher concentration of oxygen than that of oxygen-depleted blood and a lower concentration of carbon dioxide. This concentration gradient allows for gas exchange during respiration.
Partial pressure is a measure of the concentration of the individual components in a mixture of gases. The total pressure exerted by the mixture is the sum of the partial pressures of the components in the mixture. The rate of diffusion of a gas is proportional to its partial pressure within the total gas mixture. This concept is discussed further in detail below.
Lung Volumes and Capacities
Different animals have different lung capacities based on their activities. Cheetahs have evolved a much higher lung capacity than humans; it helps provide oxygen to all the muscles in the body and allows them to run very fast. Elephants also have a high lung capacity. In this case, it is not because they run fast but because they have a large body and must be able to take up oxygen in accordance with their body size.
Human lung size is determined by genetics, gender, and height. At maximal capacity, an average lung can hold almost six liters of air, but lungs do not usually operate at maximal capacity. Air in the lungs is measured in terms of lung volumes and lung capacities (Figure $1$ and Table $1$). Volume measures the amount of air for one function (such as inhalation or exhalation). Capacity is any two or more volumes (for example, how much can be inhaled from the end of a maximal exhalation).
Table $1$: Lung Volumes and Capacities (Avg Adult Male)
Volume/Capacity Definition Volume (liters) Equations
Tidal volume (TV) Amount of air inhaled during a normal breath 0.5 -
Expiratory reserve volume (ERV) Amount of air that can be exhaled after a normal exhalation 1.2 -
Inspiratory reserve volume (IRV) Amount of air that can be further inhaled after a normal inhalation 3.1 -
Residual volume (RV) Air left in the lungs after a forced exhalation 1.2 -
Vital capacity (VC) Maximum amount of air that can be moved in or out of the lungs in a single respiratory cycle 4.8 ERV+TV+IRV
Inspiratory capacity (IC) Volume of air that can be inhaled in addition to a normal exhalation 3.6 TV+IRV
Functional residual capacity (FRC) Volume of air remaining after a normal exhalation 2.4 ERV+RV
Total lung capacity (TLC) Total volume of air in the lungs after a maximal inspiration 6.0 RV+ERV+TV+IRV
Forced expiratory volume (FEV1) How much air can be forced out of the lungs over a specific time period, usually one second ~4.1 to 5.5 -
The volume in the lung can be divided into four units: tidal volume, expiratory reserve volume, inspiratory reserve volume, and residual volume. Tidal volume (TV) measures the amount of air that is inspired and expired during a normal breath. On average, this volume is around one-half liter, which is a little less than the capacity of a 20-ounce drink bottle. The expiratory reserve volume (ERV) is the additional amount of air that can be exhaled after a normal exhalation. It is the reserve amount that can be exhaled beyond what is normal. Conversely, the inspiratory reserve volume (IRV) is the additional amount of air that can be inhaled after a normal inhalation. The residual volume (RV) is the amount of air that is left after expiratory reserve volume is exhaled. The lungs are never completely empty: There is always some air left in the lungs after a maximal exhalation. If this residual volume did not exist and the lungs emptied completely, the lung tissues would stick together and the energy necessary to re-inflate the lung could be too great to overcome. Therefore, there is always some air remaining in the lungs. Residual volume is also important for preventing large fluctuations in respiratory gases (O2 and CO2). The residual volume is the only lung volume that cannot be measured directly because it is impossible to completely empty the lung of air. This volume can only be calculated rather than measured.
Capacities are measurements of two or more volumes. The vital capacity (VC) measures the maximum amount of air that can be inhaled or exhaled during a respiratory cycle. It is the sum of the expiratory reserve volume, tidal volume, and inspiratory reserve volume. The inspiratory capacity (IC) is the amount of air that can be inhaled after the end of a normal expiration. It is, therefore, the sum of the tidal volume and inspiratory reserve volume. The functional residual capacity (FRC) includes the expiratory reserve volume and the residual volume. The FRC measures the amount of additional air that can be exhaled after a normal exhalation. Lastly, the total lung capacity (TLC) is a measurement of the total amount of air that the lung can hold. It is the sum of the residual volume, expiratory reserve volume, tidal volume, and inspiratory reserve volume.
Lung volumes are measured by a technique called spirometry. An important measurement taken during spirometry is the forced expiratory volume (FEV), which measures how much air can be forced out of the lung over a specific period, usually one second (FEV1). In addition, the forced vital capacity (FVC), which is the total amount of air that can be forcibly exhaled, is measured. The ratio of these values (FEV1/FVC ratio) is used to diagnose lung diseases including asthma, emphysema, and fibrosis. If the FEV1/FVC ratio is high, the lungs are not compliant (meaning they are stiff and unable to bend properly), and the patient most likely has lung fibrosis. Patients exhale most of the lung volume very quickly. Conversely, when the FEV1/FVC ratio is low, there is resistance in the lung that is characteristic of asthma. In this instance, it is hard for the patient to get the air out of his or her lungs, and it takes a long time to reach the maximal exhalation volume. In either case, breathing is difficult and complications arise.
Career Connection: Respiratory Therapist
Respiratory therapists or respiratory practitioners evaluate and treat patients with lung and cardiovascular diseases. They work as part of a medical team to develop treatment plans for patients. Respiratory therapists may treat premature babies with underdeveloped lungs, patients with chronic conditions such as asthma, or older patients suffering from lung disease such as emphysema and chronic obstructive pulmonary disease (COPD). They may operate advanced equipment such as compressed gas delivery systems, ventilators, blood gas analyzers, and resuscitators. Specialized programs to become a respiratory therapist generally lead to a bachelor’s degree with a respiratory therapist specialty. Because of a growing aging population, career opportunities as a respiratory therapist are expected to remain strong.
Gas Pressure and Respiration
The respiratory process can be better understood by examining the properties of gases. Gases move freely, but gas particles are constantly hitting the walls of their vessel, thereby producing gas pressure.
Air is a mixture of gases, primarily nitrogen (N2; 78.6 percent), oxygen (O2; 20.9 percent), water vapor (H2O; 0.5 percent), and carbon dioxide (CO2; 0.04 percent). Each gas component of that mixture exerts a pressure. The pressure for an individual gas in the mixture is the partial pressure of that gas. Approximately 21 percent of atmospheric gas is oxygen. Carbon dioxide, however, is found in relatively small amounts, 0.04 percent. The partial pressure for oxygen is much greater than that of carbon dioxide. The partial pressure of any gas can be calculated by:
$\text{P} = \text{(P}_\text{atm}\text{)} * \text{(percent content in mixture)}\nonumber$
Patm, the atmospheric pressure, is the sum of all of the partial pressures of the atmospheric gases added together,
$\text{P}_\text{atm} = \text{P}_{\text{N}_2} + \text{P}_{\text{O}_2} + \text{P}_{\text{H}_2\text{O}} + \text{P}_{\text{CO}_2} = 760 \text{ mm Hg}\nonumber$
× (percent content in mixture).
The pressure of the atmosphere at sea level is 760 mm Hg. Therefore, the partial pressure of oxygen is:
$\text{P}_{\text{O}_2} = (760\text{ mm Hg})(0.21) = 160\text{ mm Hg}\nonumber$
and for carbon dioxide:
$\text{P}_{\text{CO}_2} = (760\text{ mm Hg})(0.0004) = 0.3\text{ mm Hg}\nonumber$
At high altitudes, Patm decreases but concentration does not change; the partial pressure decrease is due to the reduction in Patm.
When the air mixture reaches the lung, it has been humidified. The pressure of the water vapor in the lung does not change the pressure of the air, but it must be included in the partial pressure equation. For this calculation, the water pressure (47 mm Hg) is subtracted from the atmospheric pressure:
$760\text{ mm Hg} - 47\text{ mm Hg} = 713\text{ mm Hg}\nonumber$
and the partial pressure of oxygen is:
$(760\text{ mm Hg} - 47\text{ mm Hg}) (0.21) = 150\text{ mm Hg}\nonumber$
These pressures determine the gas exchange, or the flow of gas, in the system. Oxygen and carbon dioxide will flow according to their pressure gradient from high to low. Therefore, understanding the partial pressure of each gas will aid in understanding how gases move in the respiratory system.
Gas Exchange across the Alveoli
In the body, oxygen is used by cells of the body’s tissues and carbon dioxide is produced as a waste product. The ratio of carbon dioxide production to oxygen consumption is the respiratory quotient (RQ). RQ varies between 0.7 and 1.0. If just glucose were used to fuel the body, the RQ would equal one. One mole of carbon dioxide would be produced for every mole of oxygen consumed. Glucose, however, is not the only fuel for the body. Protein and fat are also used as fuels for the body. Because of this, less carbon dioxide is produced than oxygen is consumed and the RQ is, on average, about 0.7 for fat and about 0.8 for protein.
The RQ is used to calculate the partial pressure of oxygen in the alveolar spaces within the lung, the alveolar $\text{P}_{\text{O}_2}$. Above, the partial pressure of oxygen in the lungs was calculated to be 150 mm Hg. However, lungs never fully deflate with an exhalation; therefore, the inspired air mixes with this residual air and lowers the partial pressure of oxygen within the alveoli. This means that there is a lower concentration of oxygen in the lungs than is found in the air outside the body. Knowing the RQ, the partial pressure of oxygen in the alveoli can be calculated:
$\text{alveolar P}_{\text{O}_2} = \text{inspired P}_{\text{O}_2} - \frac{\text{alveolar P}_{\text{O}_2}} {\text{RQ}} \nonumber$
With an RQ of 0.8 and a $\text{P}_{\text{CO}_2}$ in the alveoli of 40 mm Hg, the alveolar $\text{P}_{\text{O}_2}$ is equal to:
$\text{alveolar P}_{\text{O}_2} = 150\text{ mm Hg} - \frac{40\text{ mm Hg}} {0.8} = \text{mm Hg}\nonumber$
Notice that this pressure is less than the external air. Therefore, the oxygen will flow from the inspired air in the lung ($\text{P}_{\text{O}_2}$ = 150 mm Hg) into the bloodstream ($\text{P}_{\text{O}_2}$ = 100 mm Hg) (Figure $2$).
In the lungs, oxygen diffuses out of the alveoli and into the capillaries surrounding the alveoli. Oxygen (about 98 percent) binds reversibly to the respiratory pigment hemoglobin found in red blood cells (RBCs). RBCs carry oxygen to the tissues where oxygen dissociates from the hemoglobin and diffuses into the cells of the tissues. More specifically, alveolar $\text{P}_{\text{O}_2}$ is higher in the alveoli $\text{P}_{\text{ALVO}_2}$ = 100 mm Hg) than blood $\text{P}_{\text{O}_2}$ (40 mm Hg) in the capillaries. Because this pressure gradient exists, oxygen diffuses down its pressure gradient, moving out of the alveoli and entering the blood of the capillaries where O2 binds to hemoglobin. At the same time, alveolar $\text{P}_{\text{CO}_2}$ is lower $\text{P}_{\text{ALVO}_2}$ = 40 mm Hg than blood $\text{P}_{\text{CO}_2}$ = (45 mm Hg). CO2 diffuses down its pressure gradient, moving out of the capillaries and entering the alveoli.
Oxygen and carbon dioxide move independently of each other; they diffuse down their own pressure gradients. As blood leaves the lungs through the pulmonary veins, the venous $\text{P}_{\text{O}_2}$= 100 mm Hg, whereas the venous $\text{P}_{\text{CO}_2}$= 40 mm Hg. As blood enters the systemic capillaries, the blood will lose oxygen and gain carbon dioxide because of the pressure difference of the tissues and blood. In systemic capillaries, $\text{P}_{\text{O}_2}$= 100 mm Hg, but in the tissue cells, $\text{P}_{\text{O}_2}$= 40 mm Hg. This pressure gradient drives the diffusion of oxygen out of the capillaries and into the tissue cells. At the same time, blood $\text{P}_{\text{CO}_2}$= 40 mm Hg and systemic tissue $\text{P}_{\text{CO}_2}$= 45 mm Hg. The pressure gradient drives CO2 out of tissue cells and into the capillaries. The blood returning to the lungs through the pulmonary arteries has a venous $\text{P}_{\text{O}_2}$= 40 mm Hg and a $\text{P}_{\text{CO}_2}$= 45 mm Hg. The blood enters the lung capillaries where the process of exchanging gases between the capillaries and alveoli begins again (Figure $2$).
Art Connection
Which of the following statements is false?
1. In the tissues, $\text{P}_{\text{O}_2}$ drops as blood passes from the arteries to the veins, while $\text{P}_{\text{CO}_2}$ increases.
2. Blood travels from the lungs to the heart to body tissues, then back to the heart, then the lungs.
3. Blood travels from the lungs to the heart to body tissues, then back to the lungs, then the heart.
4. $\text{P}_{\text{O}_2}$ is higher in air than in the lungs.
In short, the change in partial pressure from the alveoli to the capillaries drives the oxygen into the tissues and the carbon dioxide into the blood from the tissues. The blood is then transported to the lungs where differences in pressure in the alveoli result in the movement of carbon dioxide out of the blood into the lungs, and oxygen into the blood.
Link to Learning
Watch this video to learn how to carry out spirometry.
Summary
The lungs can hold a large volume of air, but they are not usually filled to maximal capacity. Lung volume measurements include tidal volume, expiratory reserve volume, inspiratory reserve volume, and residual volume. The sum of these equals the total lung capacity. Gas movement into or out of the lungs is dependent on the pressure of the gas. Air is a mixture of gases; therefore, the partial pressure of each gas can be calculated to determine how the gas will flow in the lung. The difference between the partial pressure of the gas in the air drives oxygen into the tissues and carbon dioxide out of the body.
Art Connections
Figure $2$: Which of the following statements is false?
1. In the tissues, $\text{P}_{\text{O}_2}$ drops as blood passes from the arteries to the veins, while $\text{P}_{\text{CO}_2}$ increases.
2. Blood travels from the lungs to the heart to body tissues, then back to the heart, then the lungs.
3. Blood travels from the lungs to the heart to body tissues, then back to the lungs, then the heart.
4. $\text{P}_{\text{O}_2}$ is higher in air than in the lungs.
Answer
C
Glossary
alveolar $\text{P}_{\text{O}_2}$
partial pressure of oxygen in the alveoli (usually around 100 mmHg)
expiratory reserve volume (ERV)
amount of additional air that can be exhaled after a normal exhalation
FEV1/FVC ratio
ratio of how much air can be forced out of the lung in one second to the total amount that is forced out of the lung; a measurement of lung function that can be used to detect disease states
forced expiratory volume (FEV)
(also, forced vital capacity) measure of how much air can be forced out of the lung from maximal inspiration over a specific amount of time
functional residual capacity (FRC)
expiratory reserve volume plus residual volume
inspiratory capacity (IC)
tidal volume plus inspiratory reserve volume
inspiratory reserve volume (IRV)
amount of additional air that can be inspired after a normal inhalation
lung capacity
measurement of two or more lung volumes (how much air can be inhaled from the end of an expiration to maximal capacity)
lung volume
measurement of air for one lung function (normal inhalation or exhalation)
partial pressure
amount of pressure exerted by one gas within a mixture of gases
residual volume (RV)
amount of air remaining in the lung after a maximal expiration
respiratory quotient (RQ)
ratio of carbon dioxide production to each oxygen molecule consumed
spirometry
method to measure lung volumes and to diagnose lung diseases
tidal volume (TV)
amount of air that is inspired and expired during normal breathing
total lung capacity (TLC)
sum of the residual volume, expiratory reserve volume, tidal volume, and inspiratory reserve volume
venous $\text{P}_{\text{CO}_2}$
partial pressure of carbon dioxide in the veins (40 mm Hg in the pulmonary veins)
venous $\text{P}_{\text{O}_2}$
partial pressure of oxygen in the veins (100 mm Hg in the pulmonary veins)
vital capacity (VC)
sum of the expiratory reserve volume, tidal volume, and inspiratory reserve volume
47.03: Lungs
Skills to Develop
• Describe how the structures of the lungs and thoracic cavity control the mechanics of breathing
• Explain the importance of compliance and resistance in the lungs
• Discuss problems that may arise due to a V/Q mismatch
Mammalian lungs are located in the thoracic cavity where they are surrounded and protected by the rib cage, intercostal muscles, and bound by the chest wall. The bottom of the lungs is contained by the diaphragm, a skeletal muscle that facilitates breathing. Breathing requires the coordination of the lungs, the chest wall, and most importantly, the diaphragm.
Types of Breathing
Amphibians have evolved multiple ways of breathing. Young amphibians, like tadpoles, use gills to breathe, and they don’t leave the water. Some amphibians retain gills for life. As the tadpole grows, the gills disappear and lungs grow. These lungs are primitive and not as evolved as mammalian lungs. Adult amphibians are lacking or have a reduced diaphragm, so breathing via lungs is forced. The other means of breathing for amphibians is diffusion across the skin. To aid this diffusion, amphibian skin must remain moist.
Birds face a unique challenge with respect to breathing: They fly. Flying consumes a great amount of energy; therefore, birds require a lot of oxygen to aid their metabolic processes. Birds have evolved a respiratory system that supplies them with the oxygen needed to enable flying. Similar to mammals, birds have lungs, which are organs specialized for gas exchange. Oxygenated air, taken in during inhalation, diffuses across the surface of the lungs into the bloodstream, and carbon dioxide diffuses from the blood into the lungs and expelled during exhalation. The details of breathing between birds and mammals differ substantially.
In addition to lungs, birds have air sacs inside their body. Air flows in one direction from the posterior air sacs to the lungs and out of the anterior air sacs. The flow of air is in the opposite direction from blood flow, and gas exchange takes place much more efficiently. This type of breathing enables birds to obtain the requisite oxygen, even at higher altitudes where the oxygen concentration is low. This directionality of airflow requires two cycles of air intake and exhalation to completely get the air out of the lungs.
Evolution Connection: Avian Respiration
Birds have evolved a respiratory system that enables them to fly. Flying is a high-energy process and requires a lot of oxygen. Furthermore, many birds fly in high altitudes where the concentration of oxygen in low. How did birds evolve a respiratory system that is so unique?
Decades of research by paleontologists have shown that birds evolved from therapods, meat-eating dinosaurs (Figure \(1\)). In fact, fossil evidence shows that meat-eating dinosaurs that lived more than 100 million years ago had a similar flow-through respiratory system with lungs and air sacs. Archaeopteryx and Xiaotingia, for example, were flying dinosaurs and are believed to be early precursors of birds.
Most of us consider that dinosaurs are extinct. However, modern birds are descendants of avian dinosaurs. The respiratory system of modern birds has been evolving for hundreds of millions of years.
All mammals have lungs that are the main organs for breathing. Lung capacity has evolved to support the animal’s activities. During inhalation, the lungs expand with air, and oxygen diffuses across the lung’s surface and enters the bloodstream. During exhalation, the lungs expel air and lung volume decreases. In the next few sections, the process of human breathing will be explained.
The Mechanics of Human Breathing
Boyle’s Law is the gas law that states that in a closed space, pressure and volume are inversely related. As volume decreases, pressure increases and vice versa (Figure \(2\)). The relationship between gas pressure and volume helps to explain the mechanics of breathing.
There is always a slightly negative pressure within the thoracic cavity, which aids in keeping the airways of the lungs open. During inhalation, volume increases as a result of contraction of the diaphragm, and pressure decreases (according to Boyle’s Law). This decrease of pressure in the thoracic cavity relative to the environment makes the cavity less than the atmosphere (Figure \(3\)a). Because of this drop in pressure, air rushes into the respiratory passages. To increase the volume of the lungs, the chest wall expands. This results from the contraction of the intercostal muscles, the muscles that are connected to the rib cage. Lung volume expands because the diaphragm contracts and the intercostals muscles contract, thus expanding the thoracic cavity. This increase in the volume of the thoracic cavity lowers pressure compared to the atmosphere, so air rushes into the lungs, thus increasing its volume. The resulting increase in volume is largely attributed to an increase in alveolar space, because the bronchioles and bronchi are stiff structures that do not change in size.
The chest wall expands out and away from the lungs. The lungs are elastic; therefore, when air fills the lungs, the elastic recoil within the tissues of the lung exerts pressure back toward the interior of the lungs. These outward and inward forces compete to inflate and deflate the lung with every breath. Upon exhalation, the lungs recoil to force the air out of the lungs, and the intercostal muscles relax, returning the chest wall back to its original position (Figure \(3\)b). The diaphragm also relaxes and moves higher into the thoracic cavity. This increases the pressure within the thoracic cavity relative to the environment, and air rushes out of the lungs. The movement of air out of the lungs is a passive event. No muscles are contracting to expel the air.
Each lung is surrounded by an invaginated sac. The layer of tissue that covers the lung and dips into spaces is called the visceral pleura. A second layer of parietal pleura lines the interior of the thorax (Figure \(4\)). The space between these layers, the intrapleural space, contains a small amount of fluid that protects the tissue and reduces the friction generated from rubbing the tissue layers together as the lungs contract and relax. Pleurisy results when these layers of tissue become inflamed; it is painful because the inflammation increases the pressure within the thoracic cavity and reduces the volume of the lung.
Link to Learning
View how Boyle’s Law is related to breathing and watch this video on Boyle’s Law.
The Work of Breathing
The number of breaths per minute is the respiratory rate. On average, under non-exertion conditions, the human respiratory rate is 12–15 breaths/minute. The respiratory rate contributes to the alveolar ventilation, or how much air moves into and out of the alveoli. Alveolar ventilation prevents carbon dioxide buildup in the alveoli. There are two ways to keep the alveolar ventilation constant: increase the respiratory rate while decreasing the tidal volume of air per breath (shallow breathing), or decrease the respiratory rate while increasing the tidal volume per breath. In either case, the ventilation remains the same, but the work done and type of work needed are quite different. Both tidal volume and respiratory rate are closely regulated when oxygen demand increases.
There are two types of work conducted during respiration, flow-resistive and elastic work. Flow-resistive refers to the work of the alveoli and tissues in the lung, whereas elastic work refers to the work of the intercostal muscles, chest wall, and diaphragm. Increasing the respiration rate increases the flow-resistive work of the airways and decreases the elastic work of the muscles. Decreasing the respiratory rate reverses the type of work required.
Surfactant
The air-tissue/water interface of the alveoli has a high surface tension. This surface tension is similar to the surface tension of water at the liquid-air interface of a water droplet that results in the bonding of the water molecules together. Surfactant is a complex mixture of phospholipids and lipoproteins that works to reduce the surface tension that exists between the alveoli tissue and the air found within the alveoli. By lowering the surface tension of the alveolar fluid, it reduces the tendency of alveoli to collapse.
Surfactant works like a detergent to reduce the surface tension and allows for easier inflation of the airways. When a balloon is first inflated, it takes a large amount of effort to stretch the plastic and start to inflate the balloon. If a little bit of detergent was applied to the interior of the balloon, then the amount of effort or work needed to begin to inflate the balloon would decrease, and it would become much easier to start blowing up the balloon. This same principle applies to the airways. A small amount of surfactant to the airway tissues reduces the effort or work needed to inflate those airways. Babies born prematurely sometimes do not produce enough surfactant. As a result, they suffer from respiratory distress syndrome, because it requires more effort to inflate their lungs. Surfactant is also important for preventing collapse of small alveoli relative to large alveoli.
Lung Resistance and Compliance
Pulmonary diseases reduce the rate of gas exchange into and out of the lungs. Two main causes of decreased gas exchange are compliance (how elastic the lung is) and resistance (how much obstruction exists in the airways). A change in either can dramatically alter breathing and the ability to take in oxygen and release carbon dioxide.
Examples of restrictive diseases are respiratory distress syndrome and pulmonary fibrosis. In both diseases, the airways are less compliant and they are stiff or fibrotic. There is a decrease in compliance because the lung tissue cannot bend and move. In these types of restrictive diseases, the intrapleural pressure is more positive and the airways collapse upon exhalation, which traps air in the lungs. Forced or functional vital capacity (FVC), which is the amount of air that can be forcibly exhaled after taking the deepest breath possible, is much lower than in normal patients, and the time it takes to exhale most of the air is greatly prolonged (Figure \(5\)). A patient suffering from these diseases cannot exhale the normal amount of air.
Obstructive diseases and conditions include emphysema, asthma, and pulmonary edema. In emphysema, which mostly arises from smoking tobacco, the walls of the alveoli are destroyed, decreasing the surface area for gas exchange. The overall compliance of the lungs is increased, because as the alveolar walls are damaged, lung elastic recoil decreases due to a loss of elastic fibers, and more air is trapped in the lungs at the end of exhalation. Asthma is a disease in which inflammation is triggered by environmental factors. Inflammation obstructs the airways. The obstruction may be due to edema (fluid accumulation), smooth muscle spasms in the walls of the bronchioles, increased mucus secretion, damage to the epithelia of the airways, or a combination of these events. Those with asthma or edema experience increased occlusion from increased inflammation of the airways. This tends to block the airways, preventing the proper movement of gases (Figure \(5\)). Those with obstructive diseases have large volumes of air trapped after exhalation and breathe at a very high lung volume to compensate for the lack of airway recruitment.
Dead Space: V/Q Mismatch
Pulmonary circulation pressure is very low compared to that of the systemic circulation. It is also independent of cardiac output. This is because of a phenomenon called recruitment, which is the process of opening airways that normally remain closed when cardiac output increases. As cardiac output increases, the number of capillaries and arteries that are perfused (filled with blood) increases. These capillaries and arteries are not always in use but are ready if needed. At times, however, there is a mismatch between the amount of air (ventilation, V) and the amount of blood (perfusion, Q) in the lungs. This is referred to as ventilation/perfusion (V/Q) mismatch.
There are two types of V/Q mismatch. Both produce dead space, regions of broken down or blocked lung tissue. Dead spaces can severely impact breathing, because they reduce the surface area available for gas diffusion. As a result, the amount of oxygen in the blood decreases, whereas the carbon dioxide level increases. Dead space is created when no ventilation and/or perfusion takes place. Anatomical dead space or anatomical shunt, arises from an anatomical failure, while physiological dead space or physiological shunt, arises from a functional impairment of the lung or arteries.
An example of an anatomical shunt is the effect of gravity on the lungs. The lung is particularly susceptible to changes in the magnitude and direction of gravitational forces. When someone is standing or sitting upright, the pleural pressure gradient leads to increased ventilation further down in the lung. As a result, the intrapleural pressure is more negative at the base of the lung than at the top, and more air fills the bottom of the lung than the top. Likewise, it takes less energy to pump blood to the bottom of the lung than to the top when in a prone position. Perfusion of the lung is not uniform while standing or sitting. This is a result of hydrostatic forces combined with the effect of airway pressure. An anatomical shunt develops because the ventilation of the airways does not match the perfusion of the arteries surrounding those airways. As a result, the rate of gas exchange is reduced. Note that this does not occur when lying down, because in this position, gravity does not preferentially pull the bottom of the lung down.
A physiological shunt can develop if there is infection or edema in the lung that obstructs an area. This will decrease ventilation but not affect perfusion; therefore, the V/Q ratio changes and gas exchange is affected.
The lung can compensate for these mismatches in ventilation and perfusion. If ventilation is greater than perfusion, the arterioles dilate and the bronchioles constrict. This increases perfusion and reduces ventilation. Likewise, if ventilation is less than perfusion, the arterioles constrict and the bronchioles dilate to correct the imbalance.
Link to Learning
Visit this site to view the mechanics of breathing.
Summary
The structure of the lungs and thoracic cavity control the mechanics of breathing. Upon inspiration, the diaphragm contracts and lowers. The intercostal muscles contract and expand the chest wall outward. The intrapleural pressure drops, the lungs expand, and air is drawn into the airways. When exhaling, the intercostal muscles and diaphragm relax, returning the intrapleural pressure back to the resting state. The lungs recoil and airways close. The air passively exits the lung. There is high surface tension at the air-airway interface in the lung. Surfactant, a mixture of phospholipids and lipoproteins, acts like a detergent in the airways to reduce surface tension and allow for opening of the alveoli.
Breathing and gas exchange are both altered by changes in the compliance and resistance of the lung. If the compliance of the lung decreases, as occurs in restrictive diseases like fibrosis, the airways stiffen and collapse upon exhalation. Air becomes trapped in the lungs, making breathing more difficult. If resistance increases, as happens with asthma or emphysema, the airways become obstructed, trapping air in the lungs and causing breathing to become difficult. Alterations in the ventilation of the airways or perfusion of the arteries can affect gas exchange. These changes in ventilation and perfusion, called V/Q mismatch, can arise from anatomical or physiological changes.
Glossary
alveolar ventilation
how much air is in the alveoli
anatomical dead space
(also, anatomical shunt) region of the lung that lacks proper ventilation/perfusion due to an anatomical block
compliance
measurement of the elasticity of the lung
dead space
area in the lung that lacks proper ventilation or perfusion
elastic recoil
property of the lung that drives the lung tissue inward
elastic work
work conducted by the intercostal muscles, chest wall, and diaphragm
flow-resistive
work of breathing performed by the alveoli and tissues in the lung
functional vital capacity (FVC)
amount of air that can be forcibly exhaled after taking the deepest breath possible
intercostal muscle
muscle connected to the rib cage that contracts upon inspiration
intrapleural space
space between the layers of pleura
obstructive disease
disease (such as emphysema and asthma) that arises from obstruction of the airways; compliance increases in these diseases
physiological dead space
(also, physiological shunt) region of the lung that lacks proper ventilation/perfusion due to a physiological change in the lung (like inflammation or edema)
pleura
tissue layer that surrounds the lungs and lines the interior of the thoracic cavity
pleurisy
painful inflammation of the pleural tissue layers
recruitment
process of opening airways that normally remain closed when the cardiac output increases
resistance
measurement of lung obstruction
respiratory distress syndrome
disease that arises from a deficient amount of surfactant
respiratory rate
number of breaths per minute
restrictive disease
disease that results from a restriction and decreased compliance of the alveoli; respiratory distress syndrome and pulmonary fibrosis are examples
surfactant
detergent-like liquid in the airways that lowers the surface tension of the alveoli to allow for expansion
ventilation/perfusion (V/Q) mismatch
region of the lung that lacks proper alveolar ventilation (V) and/or arterial perfusion (Q) | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/47%3A_The_Respiratory_System/47.03%3A_Lungs/47.3.01%3A_Breathing.txt |
Skills to Develop
• Describe how oxygen is bound to hemoglobin and transported to body tissues
• Explain how carbon dioxide is transported from body tissues to the lungs
Once the oxygen diffuses across the alveoli, it enters the bloodstream and is transported to the tissues where it is unloaded, and carbon dioxide diffuses out of the blood and into the alveoli to be expelled from the body. Although gas exchange is a continuous process, the oxygen and carbon dioxide are transported by different mechanisms.
Transport of Oxygen in the Blood
Although oxygen dissolves in blood, only a small amount of oxygen is transported this way. Only 1.5 percent of oxygen in the blood is dissolved directly into the blood itself. Most oxygen—98.5 percent—is bound to a protein called hemoglobin and carried to the tissues.
Hemoglobin
Hemoglobin, or Hb, is a protein molecule found in red blood cells (erythrocytes) made of four subunits: two alpha subunits and two beta subunits (Figure $1$). Each subunit surrounds a central heme group that contains iron and binds one oxygen molecule, allowing each hemoglobin molecule to bind four oxygen molecules. Molecules with more oxygen bound to the heme groups are brighter red. As a result, oxygenated arterial blood where the Hb is carrying four oxygen molecules is bright red, while venous blood that is deoxygenated is darker red.
It is easier to bind a second and third oxygen molecule to Hb than the first molecule. This is because the hemoglobin molecule changes its shape, or conformation, as oxygen binds. The fourth oxygen is then more difficult to bind. The binding of oxygen to hemoglobin can be plotted as a function of the partial pressure of oxygen in the blood (x-axis) versus the relative Hb-oxygen saturation (y-axis). The resulting graph—an oxygen dissociation curve—is sigmoidal, or S-shaped (Figure $2$). As the partial pressure of oxygen increases, the hemoglobin becomes increasingly saturated with oxygen.
Art Connection
The kidneys are responsible for removing excess H+ ions from the blood. If the kidneys fail, what would happen to blood pH and to hemoglobin affinity for oxygen?
Factors That Affect Oxygen Binding
The oxygen-carrying capacity of hemoglobin determines how much oxygen is carried in the blood. In addition to $\text{P}_{\text{O}_2}$, other environmental factors and diseases can affect oxygen carrying capacity and delivery.
Carbon dioxide levels, blood pH, and body temperature affect oxygen-carrying capacity (Figure $2$). When carbon dioxide is in the blood, it reacts with water to form bicarbonate ($\text{HCO}_3^-$) and hydrogen ions (H+). As the level of carbon dioxide in the blood increases, more H+ is produced and the pH decreases. This increase in carbon dioxide and subsequent decrease in pH reduce the affinity of hemoglobin for oxygen. The oxygen dissociates from the Hb molecule, shifting the oxygen dissociation curve to the right. Therefore, more oxygen is needed to reach the same hemoglobin saturation level as when the pH was higher. A similar shift in the curve also results from an increase in body temperature. Increased temperature, such as from increased activity of skeletal muscle, causes the affinity of hemoglobin for oxygen to be reduced.
Diseases like sickle cell anemia and thalassemia decrease the blood’s ability to deliver oxygen to tissues and its oxygen-carrying capacity. In sickle cell anemia, the shape of the red blood cell is crescent-shaped, elongated, and stiffened, reducing its ability to deliver oxygen (Figure $3$). In this form, red blood cells cannot pass through the capillaries. This is painful when it occurs. Thalassemia is a rare genetic disease caused by a defect in either the alpha or the beta subunit of Hb. Patients with thalassemia produce a high number of red blood cells, but these cells have lower-than-normal levels of hemoglobin. Therefore, the oxygen-carrying capacity is diminished.
Transport of Carbon Dioxide in the Blood
Carbon dioxide molecules are transported in the blood from body tissues to the lungs by one of three methods: dissolution directly into the blood, binding to hemoglobin, or carried as a bicarbonate ion. Several properties of carbon dioxide in the blood affect its transport. First, carbon dioxide is more soluble in blood than oxygen. About 5 to 7 percent of all carbon dioxide is dissolved in the plasma. Second, carbon dioxide can bind to plasma proteins or can enter red blood cells and bind to hemoglobin. This form transports about 10 percent of the carbon dioxide. When carbon dioxide binds to hemoglobin, a molecule called carbaminohemoglobin is formed. Binding of carbon dioxide to hemoglobin is reversible. Therefore, when it reaches the lungs, the carbon dioxide can freely dissociate from the hemoglobin and be expelled from the body.
Third, the majority of carbon dioxide molecules (85 percent) are carried as part of the bicarbonate buffer system. In this system, carbon dioxide diffuses into the red blood cells. Carbonic anhydrase (CA) within the red blood cells quickly converts the carbon dioxide into carbonic acid (H2CO3). Carbonic acid is an unstable intermediate molecule that immediately dissociates into bicarbonate ions ($\text{HCO}_3^-$) and hydrogen (H+) ions. Since carbon dioxide is quickly converted into bicarbonate ions, this reaction allows for the continued uptake of carbon dioxide into the blood down its concentration gradient. It also results in the production of H+ ions. If too much H+ is produced, it can alter blood pH. However, hemoglobin binds to the free H+ ions and thus limits shifts in pH. The newly synthesized bicarbonate ion is transported out of the red blood cell into the liquid component of the blood in exchange for a chloride ion (Cl-); this is called the chloride shift. When the blood reaches the lungs, the bicarbonate ion is transported back into the red blood cell in exchange for the chloride ion. The H+ ion dissociates from the hemoglobin and binds to the bicarbonate ion. This produces the carbonic acid intermediate, which is converted back into carbon dioxide through the enzymatic action of CA. The carbon dioxide produced is expelled through the lungs during exhalation.
$\text{CO}_2 + \text{H}_2\text{O} \leftrightharpoons \underset{\text{(carbonic acid)}}{\text{H}_2\text{CO}_3} \leftrightarrow \underset{\text{(bicarbonate)}}{\text{HCO}_3 + \text{H}^+}\nonumber$
The benefit of the bicarbonate buffer system is that carbon dioxide is “soaked up” into the blood with little change to the pH of the system. This is important because it takes only a small change in the overall pH of the body for severe injury or death to result. The presence of this bicarbonate buffer system also allows for people to travel and live at high altitudes: When the partial pressure of oxygen and carbon dioxide change at high altitudes, the bicarbonate buffer system adjusts to regulate carbon dioxide while maintaining the correct pH in the body.
Carbon Monoxide Poisoning
While carbon dioxide can readily associate and dissociate from hemoglobin, other molecules such as carbon monoxide (CO) cannot. Carbon monoxide has a greater affinity for hemoglobin than oxygen. Therefore, when carbon monoxide is present, it binds to hemoglobin preferentially over oxygen. As a result, oxygen cannot bind to hemoglobin, so very little oxygen is transported through the body (Figure $4$). Carbon monoxide is a colorless, odorless gas and is therefore difficult to detect. It is produced by gas-powered vehicles and tools. Carbon monoxide can cause headaches, confusion, and nausea; long-term exposure can cause brain damage or death. Administering 100 percent (pure) oxygen is the usual treatment for carbon monoxide poisoning. Administration of pure oxygen speeds up the separation of carbon monoxide from hemoglobin.
Summary
Hemoglobin is a protein found in red blood cells that is comprised of two alpha and two beta subunits that surround an iron-containing heme group. Oxygen readily binds this heme group. The ability of oxygen to bind increases as more oxygen molecules are bound to heme. Disease states and altered conditions in the body can affect the binding ability of oxygen, and increase or decrease its ability to dissociate from hemoglobin.
Carbon dioxide can be transported through the blood via three methods. It is dissolved directly in the blood, bound to plasma proteins or hemoglobin, or converted into bicarbonate. The majority of carbon dioxide is transported as part of the bicarbonate system. Carbon dioxide diffuses into red blood cells. Inside, carbonic anhydrase converts carbon dioxide into carbonic acid (H2CO3), which is subsequently hydrolyzed into bicarbonate ($\text{HCO}_3^-$) and H+. The H+ ion binds to hemoglobin in red blood cells, and bicarbonate is transported out of the red blood cells in exchange for a chloride ion. This is called the chloride shift. Bicarbonate leaves the red blood cells and enters the blood plasma. In the lungs, bicarbonate is transported back into the red blood cells in exchange for chloride. The H+ dissociates from hemoglobin and combines with bicarbonate to form carbonic acid with the help of carbonic anhydrase, which further catalyzes the reaction to convert carbonic acid back into carbon dioxide and water. The carbon dioxide is then expelled from the lungs.
Art Connections
Figure $2$: The kidneys are responsible for removing excess H+ ions from the blood. If the kidneys fail, what would happen to blood pH and to hemoglobin affinity for oxygen?
Answer
The blood pH will drop and hemoglobin affinity for oxygen will decrease.
Glossary
bicarbonate buffer system
system in the blood that absorbs carbon dioxide and regulates pH levels
bicarbonate ($\text{HCO}_3^-$) ion
ion created when carbonic acid dissociates into H+ and $(\ce{HCO3-})$
carbaminohemoglobin
molecule that forms when carbon dioxide binds to hemoglobin
carbonic anhydrase (CA)
enzyme that catalyzes carbon dioxide and water into carbonic acid
chloride shift
chloride shift exchange of chloride for bicarbonate into or out of the red blood cell
heme group
centralized iron-containing group that is surrounded by the alpha and beta subunits of hemoglobin
hemoglobin
molecule in red blood cells that can bind oxygen, carbon dioxide, and carbon monoxide
oxygen-carrying capacity
amount of oxygen that can be transported in the blood
oxygen dissociation curve
curve depicting the affinity of oxygen for hemoglobin
sickle cell anemia
genetic disorder that affects the shape of red blood cells, and their ability to transport oxygen and move through capillaries
thalassemia
rare genetic disorder that results in mutation of the alpha or beta subunits of hemoglobin, creating smaller red blood cells with less hemoglobin | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/47%3A_The_Respiratory_System/47.05%3A_Transport_of_Gases_in_Body_Fluid.txt |
• 48.1: Invertebrate Circulatory Systems
In all animals, except a few simple types, the circulatory system is used to transport nutrients and gases through the body. Simple diffusion allows some water, nutrient, waste, and gas exchange into primitive animals that are only a few cell layers thick; however, bulk flow is the only method by which the entire body of larger more complex organisms is accessed.
• 48.2: Components of Vertebrate Blood
Blood is the liquid that moves through the vessels and includes plasma (the liquid portion, which contains water, proteins, salts, lipids, and glucose) and the cells (red and white cells) and cell fragments called platelets. Blood plasma is actually the dominant component of blood and contains the water, proteins, electrolytes, lipids, and glucose. The cells are responsible for carrying the gases (red cells) and immune the response (white). The platelets are responsible for blood clotting.
• 48.3: Vertebrate Circulatory System
The heart is a complex muscle that pumps blood through the three divisions of the circulatory system: the coronary (vessels that serve the heart), pulmonary (heart and lungs), and systemic (systems of the body). Coronary circulation intrinsic to the heart takes blood directly from the main artery (aorta) coming from the heart.
• 48.4: Cardiac Cycle, Electrical Conduction, ECG, and Cardiac Output
Blood pressure is the pressure exerted by blood on the walls of a blood vessel that helps to push blood through the body. Systolic blood pressure measures the amount of pressure that blood exerts on vessels while the heart is beating. The optimal systolic blood pressure is 120 mmHg. Diastolic blood pressure measures the pressure in the vessels between heartbeats. The optimal diastolic blood pressure is 80 mmHg.
• 48.5: Blood Pressure and Blood Vessels
48: The Circulatory System
Skills to Develop
• Describe an open and closed circulatory system
• Describe interstitial fluid and hemolymph
• Compare and contrast the organization and evolution of the vertebrate circulatory system.
In all animals, except a few simple types, the circulatory system is used to transport nutrients and gases through the body. Simple diffusion allows some water, nutrient, waste, and gas exchange into primitive animals that are only a few cell layers thick; however, bulk flow is the only method by which the entire body of larger more complex organisms is accessed.
Circulatory System Architecture
The circulatory system is effectively a network of cylindrical vessels: the arteries, veins, and capillaries that emanate from a pump, the heart. In all vertebrate organisms, as well as some invertebrates, this is a closed-loop system, in which the blood is not free in a cavity. In a closed circulatory system, blood is contained inside blood vessels and circulates unidirectionally from the heart around the systemic circulatory route, then returns to the heart again, as illustrated in Figure \(1\)a. As opposed to a closed system, arthropods—including insects, crustaceans, and most mollusks—have an open circulatory system, as illustrated in Figure \(1\)b. In an open circulatory system, the blood is not enclosed in the blood vessels but is pumped into a cavity called a hemocoel and is called hemolymph because the blood mixes with the interstitial fluid. As the heart beats and the animal moves, the hemolymph circulates around the organs within the body cavity and then reenters the hearts through openings called ostia. This movement allows for gas and nutrient exchange. An open circulatory system does not use as much energy as a closed system to operate or to maintain; however, there is a trade-off with the amount of blood that can be moved to metabolically active organs and tissues that require high levels of oxygen. In fact, one reason that insects with wing spans of up to two feet wide (70 cm) are not around today is probably because they were outcompeted by the arrival of birds 150 million years ago. Birds, having a closed circulatory system, are thought to have moved more agilely, allowing them to get food faster and possibly to prey on the insects.
Circulatory System Variation in Animals
The circulatory system varies from simple systems in invertebrates to more complex systems in vertebrates. The simplest animals, such as the sponges (Porifera) and rotifers (Rotifera), do not need a circulatory system because diffusion allows adequate exchange of water, nutrients, and waste, as well as dissolved gases, as shown in Figure \(2\)a. Organisms that are more complex but still only have two layers of cells in their body plan, such as jellies (Cnidaria) and comb jellies (Ctenophora) also use diffusion through their epidermis and internally through the gastrovascular compartment. Both their internal and external tissues are bathed in an aqueous environment and exchange fluids by diffusion on both sides, as illustrated in Figure \(2\)b. Exchange of fluids is assisted by the pulsing of the jellyfish body.
For more complex organisms, diffusion is not efficient for cycling gases, nutrients, and waste effectively through the body; therefore, more complex circulatory systems evolved. Most arthropods and many mollusks have open circulatory systems. In an open system, an elongated beating heart pushes the hemolymph through the body and muscle contractions help to move fluids. The larger more complex crustaceans, including lobsters, have developed arterial-like vessels to push blood through their bodies, and the most active mollusks, such as squids, have evolved a closed circulatory system and are able to move rapidly to catch prey. Closed circulatory systems are a characteristic of vertebrates; however, there are significant differences in the structure of the heart and the circulation of blood between the different vertebrate groups due to adaptation during evolution and associated differences in anatomy. Figure \(3\) illustrates the basic circulatory systems of some vertebrates: fish, amphibians, reptiles, and mammals.
As illustrated in Figure \(3\)a Fish have a single circuit for blood flow and a two-chambered heart that has only a single atrium and a single ventricle. The atrium collects blood that has returned from the body and the ventricle pumps the blood to the gills where gas exchange occurs and the blood is re-oxygenated; this is called gill circulation. The blood then continues through the rest of the body before arriving back at the atrium; this is called systemic circulation. This unidirectional flow of blood produces a gradient of oxygenated to deoxygenated blood around the fish’s systemic circuit. The result is a limit in the amount of oxygen that can reach some of the organs and tissues of the body, reducing the overall metabolic capacity of fish.
In amphibians, reptiles, birds, and mammals, blood flow is directed in two circuits: one through the lungs and back to the heart, which is called pulmonary circulation, and the other throughout the rest of the body and its organs including the brain (systemic circulation). In amphibians, gas exchange also occurs through the skin during pulmonary circulation and is referred to as pulmocutaneous circulation.
As shown in Figure \(3\)b, amphibians have a three-chambered heart that has two atria and one ventricle rather than the two-chambered heart of fish. The two atria (superior heart chambers) receive blood from the two different circuits (the lungs and the systems), and then there is some mixing of the blood in the heart’s ventricle (inferior heart chamber), which reduces the efficiency of oxygenation. The advantage to this arrangement is that high pressure in the vessels pushes blood to the lungs and body. The mixing is mitigated by a ridge within the ventricle that diverts oxygen-rich blood through the systemic circulatory system and deoxygenated blood to the pulmocutaneous circuit. For this reason, amphibians are often described as having double circulation.
Most reptiles also have a three-chambered heart similar to the amphibian heart that directs blood to the pulmonary and systemic circuits, as shown in Figure \(3\)c. The ventricle is divided more effectively by a partial septum, which results in less mixing of oxygenated and deoxygenated blood. Some reptiles (alligators and crocodiles) are the most primitive animals to exhibit a four-chambered heart. Crocodilians have a unique circulatory mechanism where the heart shunts blood from the lungs toward the stomach and other organs during long periods of submergence, for instance, while the animal waits for prey or stays underwater waiting for prey to rot. One adaptation includes two main arteries that leave the same part of the heart: one takes blood to the lungs and the other provides an alternate route to the stomach and other parts of the body. Two other adaptations include a hole in the heart between the two ventricles, called the foramen of Panizza, which allows blood to move from one side of the heart to the other, and specialized connective tissue that slows the blood flow to the lungs. Together these adaptations have made crocodiles and alligators one of the most evolutionarily successful animal groups on earth.
In mammals and birds, the heart is also divided into four chambers: two atria and two ventricles, as illustrated in Figure 40.1.3d. The oxygenated blood is separated from the deoxygenated blood, which improves the efficiency of double circulation and is probably required for the warm-blooded lifestyle of mammals and birds. The four-chambered heart of birds and mammals evolved independently from a three-chambered heart. The independent evolution of the same or a similar biological trait is referred to as convergent evolution.
Summary
In most animals, the circulatory system is used to transport blood through the body. Some primitive animals use diffusion for the exchange of water, nutrients, and gases. However, complex organisms use the circulatory system to carry gases, nutrients, and waste through the body. Circulatory systems may be open (mixed with the interstitial fluid) or closed (separated from the interstitial fluid). Closed circulatory systems are a characteristic of vertebrates; however, there are significant differences in the structure of the heart and the circulation of blood between the different vertebrate groups due to adaptions during evolution and associated differences in anatomy. Fish have a two-chambered heart with unidirectional circulation. Amphibians have a three-chambered heart, which has some mixing of the blood, and they have double circulation. Most non-avian reptiles have a three-chambered heart, but have little mixing of the blood; they have double circulation. Mammals and birds have a four-chambered heart with no mixing of the blood and double circulation.
Glossary
atrium
(plural: atria) chamber of the heart that receives blood from the veins and sends blood to the ventricles
closed circulatory system
system in which the blood is separated from the bodily interstitial fluid and contained in blood vessels
double circulation
flow of blood in two circuits: the pulmonary circuit through the lungs and the systemic circuit through the organs and body
gill circulation
circulatory system that is specific to animals with gills for gas exchange; the blood flows through the gills for oxygenation
hemocoel
cavity into which blood is pumped in an open circulatory system
hemolymph
mixture of blood and interstitial fluid that is found in insects and other arthropods as well as most mollusks
interstitial fluid
fluid between cells
open circulatory system
system in which the blood is mixed with interstitial fluid and directly covers the organs
ostium
(plural: ostia) holes between blood vessels that allow the movement of hemolymph through the body of insects, arthropods, and mollusks with open circulatory systems
pulmocutaneous circulation
circulatory system in amphibians; the flow of blood to the lungs and the moist skin for gas exchange
pulmonary circulation
flow of blood away from the heart through the lungs where oxygenation occurs and then returns to the heart again
systemic circulation
flow of blood away from the heart to the brain, liver, kidneys, stomach, and other organs, the limbs, and the muscles of the body, and then the return of this blood to the heart
unidirectional circulation
flow of blood in a single circuit; occurs in fish where the blood flows through the gills, then past the organs and the rest of the body, before returning to the heart
ventricle
(heart) large inferior chamber of the heart that pumps blood into arteries | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/48%3A_The_Circulatory_System/48.01%3A_Invertebrate_Circulatory_Systems.txt |
Skills to Develop
• List the basic components of the blood
• Compare red and white blood cells
• Describe blood plasma and serum
Hemoglobin is responsible for distributing oxygen, and to a lesser extent, carbon dioxide, throughout the circulatory systems of humans, vertebrates, and many invertebrates. The blood is more than the proteins, though. Blood is actually a term used to describe the liquid that moves through the vessels and includes plasma (the liquid portion, which contains water, proteins, salts, lipids, and glucose) and the cells (red and white cells) and cell fragments called platelets. Blood plasma is actually the dominant component of blood and contains the water, proteins, electrolytes, lipids, and glucose. The cells are responsible for carrying the gases (red cells) and immune the response (white). The platelets are responsible for blood clotting. Interstitial fluid that surrounds cells is separate from the blood, but in hemolymph, they are combined. In humans, cellular components make up approximately 45 percent of the blood and the liquid plasma 55 percent. Blood is 20 percent of a person’s extracellular fluid and eight percent of weight.
The Role of Blood in the Body
Blood, like the human blood illustrated in Figure \(1\) is important for regulation of the body’s systems and homeostasis. Blood helps maintain homeostasis by stabilizing pH, temperature, osmotic pressure, and by eliminating excess heat. Blood supports growth by distributing nutrients and hormones, and by removing waste. Blood plays a protective role by transporting clotting factors and platelets to prevent blood loss and transporting the disease-fighting agents or white blood cells to sites of infection.
Red Blood Cells
Red blood cells, or erythrocytes (erythro- = “red”; -cyte = “cell”), are specialized cells that circulate through the body delivering oxygen to cells; they are formed from stem cells in the bone marrow. In mammals, red blood cells are small biconcave cells that at maturity do not contain a nucleus or mitochondria and are only 7–8 µm in size. In birds and non-avian reptiles, a nucleus is still maintained in red blood cells.
The red coloring of blood comes from the iron-containing protein hemoglobin, illustrated in Figure \(2\)a. The principal job of this protein is to carry oxygen, but it also transports carbon dioxide as well. Hemoglobin is packed into red blood cells at a rate of about 250 million molecules of hemoglobin per cell. Each hemoglobin molecule binds four oxygen molecules so that each red blood cell carries one billion molecules of oxygen. There are approximately 25 trillion red blood cells in the five liters of blood in the human body, which could carry up to 25 sextillion (25 × 1021) molecules of oxygen in the body at any time. In mammals, the lack of organelles in erythrocytes leaves more room for the hemoglobin molecules, and the lack of mitochondria also prevents use of the oxygen for metabolic respiration. Only mammals have anucleated red blood cells, and some mammals (camels, for instance) even have nucleated red blood cells. The advantage of nucleated red blood cells is that these cells can undergo mitosis. Anucleated red blood cells metabolize anaerobically (without oxygen), making use of a primitive metabolic pathway to produce ATP and increase the efficiency of oxygen transport.
Not all organisms use hemoglobin as the method of oxygen transport. Invertebrates that utilize hemolymph rather than blood use different pigments to bind to the oxygen. These pigments use copper or iron to the oxygen. Invertebrates have a variety of other respiratory pigments. Hemocyanin, a blue-green, copper-containing protein, illustrated in Figure \(2\)b is found in mollusks, crustaceans, and some of the arthropods. Chlorocruorin, a green-colored, iron-containing pigment is found in four families of polychaete tubeworms. Hemerythrin, a red, iron-containing protein is found in some polychaete worms and annelids and is illustrated in Figure \(2\)c. Despite the name, hemerythrin does not contain a heme group and its oxygen-carrying capacity is poor compared to hemoglobin.
The small size and large surface area of red blood cells allows for rapid diffusion of oxygen and carbon dioxide across the plasma membrane. In the lungs, carbon dioxide is released and oxygen is taken in by the blood. In the tissues, oxygen is released from the blood and carbon dioxide is bound for transport back to the lungs. Studies have found that hemoglobin also binds nitrous oxide (NO). NO is a vasodilator that relaxes the blood vessels and capillaries and may help with gas exchange and the passage of red blood cells through narrow vessels. Nitroglycerin, a heart medication for angina and heart attacks, is converted to NO to help relax the blood vessels and increase oxygen flow through the body.
A characteristic of red blood cells is their glycolipid and glycoprotein coating; these are lipids and proteins that have carbohydrate molecules attached. In humans, the surface glycoproteins and glycolipids on red blood cells vary between individuals, producing the different blood types, such as A, B, and O. Red blood cells have an average life span of 120 days, at which time they are broken down and recycled in the liver and spleen by phagocytic macrophages, a type of white blood cell.
White Blood Cells
White blood cells, also called leukocytes (leuko = white), make up approximately one percent by volume of the cells in blood. The role of white blood cells is very different than that of red blood cells: they are primarily involved in the immune response to identify and target pathogens, such as invading bacteria, viruses, and other foreign organisms. White blood cells are formed continually; some only live for hours or days, but some live for years.
The morphology of white blood cells differs significantly from red blood cells. They have nuclei and do not contain hemoglobin. The different types of white blood cells are identified by their microscopic appearance after histologic staining, and each has a different specialized function. The two main groups, both illustrated in Figure \(3\) are the granulocytes, which include the neutrophils, eosinophils, and basophils, and the agranulocytes, which include the monocytes and lymphocytes.
Granulocytes contain granules in their cytoplasm; the agranulocytes are so named because of the lack of granules in their cytoplasm. Some leukocytes become macrophages that either stay at the same site or move through the blood stream and gather at sites of infection or inflammation where they are attracted by chemical signals from foreign particles and damaged cells. Lymphocytes are the primary cells of the immune system and include B cells, T cells, and natural killer cells. B cells destroy bacteria and inactivate their toxins. They also produce antibodies. T cells attack viruses, fungi, some bacteria, transplanted cells, and cancer cells. T cells attack viruses by releasing toxins that kill the viruses. Natural killer cells attack a variety of infectious microbes and certain tumor cells.
One reason that HIV poses significant management challenges is because the virus directly targets T cells by gaining entry through a receptor. Once inside the cell, HIV then multiplies using the T cell’s own genetic machinery. After the HIV virus replicates, it is transmitted directly from the infected T cell to macrophages. The presence of HIV can remain unrecognized for an extensive period of time before full disease symptoms develop.
Platelets and Coagulation Factors
Blood must clot to heal wounds and prevent excess blood loss. Small cell fragments called platelets (thrombocytes) are attracted to the wound site where they adhere by extending many projections and releasing their contents. These contents activate other platelets and also interact with other coagulation factors, which convert fibrinogen, a water-soluble protein present in blood serum into fibrin (a non-water soluble protein), causing the blood to clot. Many of the clotting factors require vitamin K to work, and vitamin K deficiency can lead to problems with blood clotting. Many platelets converge and stick together at the wound site forming a platelet plug (also called a fibrin clot), as illustrated in Figure \(4\)b. The plug or clot lasts for a number of days and stops the loss of blood. Platelets are formed from the disintegration of larger cells called megakaryocytes, like that shown in Figure \(4\)a. For each megakaryocyte, 2000–3000 platelets are formed with 150,000 to 400,000 platelets present in each cubic millimeter of blood. Each platelet is disc shaped and 2–4 μm in diameter. They contain many small vesicles but do not contain a nucleus.
Plasma and Serum
The liquid component of blood is called plasma, and it is separated by spinning or centrifuging the blood at high rotations (3000 rpm or higher). The blood cells and platelets are separated by centrifugal forces to the bottom of a specimen tube. The upper liquid layer, the plasma, consists of 90 percent water along with various substances required for maintaining the body’s pH, osmotic load, and for protecting the body. The plasma also contains the coagulation factors and antibodies.
The plasma component of blood without the coagulation factors is called the serum. Serum is similar to interstitial fluid in which the correct composition of key ions acting as electrolytes is essential for normal functioning of muscles and nerves. Other components in the serum include proteins that assist with maintaining pH and osmotic balance while giving viscosity to the blood. The serum also contains antibodies, specialized proteins that are important for defense against viruses and bacteria. Lipids, including cholesterol, are also transported in the serum, along with various other substances including nutrients, hormones, metabolic waste, plus external substances, such as, drugs, viruses, and bacteria.
Human serum albumin is the most abundant protein in human blood plasma and is synthesized in the liver. Albumin, which constitutes about half of the blood serum protein, transports hormones and fatty acids, buffers pH, and maintains osmotic pressures. Immunoglobin is a protein antibody produced in the mucosal lining and plays an important role in antibody mediated immunity.
Evolution Connection: Blood Types
Related to Proteins on the Surface of the Red Blood Cells Red blood cells are coated in antigens made of glycolipids and glycoproteins. The composition of these molecules is determined by genetics, which have evolved over time. In humans, the different surface antigens are grouped into 24 different blood groups with more than 100 different antigens on each red blood cell. The two most well known blood groups are the ABO, shown in Figure \(5\), and Rh systems. The surface antigens in the ABO blood group are glycolipids, called antigen A and antigen B. People with blood type A have antigen A, those with blood type B have antigen B, those with blood type AB have both antigens, and people with blood type O have neither antigen. Antibodies called agglutinougens are found in the blood plasma and react with the A or B antigens, if the two are mixed. When type A and type B blood are combined, agglutination (clumping) of the blood occurs because of antibodies in the plasma that bind with the opposing antigen; this causes clots that coagulate in the kidney causing kidney failure. Type O blood has neither A or B antigens, and therefore, type O blood can be given to all blood types. Type O negative blood is the universal donor. Type AB positive blood is the universal acceptor because it has both A and B antigen. The ABO blood groups were discovered in 1900 and 1901 by Karl Landsteiner at the University of Vienna.
The Rh blood group was first discovered in Rhesus monkeys. Most people have the Rh antigen (Rh+) and do not have anti-Rh antibodies in their blood. The few people who do not have the Rh antigen and are Rh– can develop anti-Rh antibodies if exposed to Rh+ blood. This can happen after a blood transfusion or after an Rh– woman has an Rh+ baby. The first exposure does not usually cause a reaction; however, at the second exposure, enough antibodies have built up in the blood to produce a reaction that causes agglutination and breakdown of red blood cells. An injection can prevent this reaction.
Summary
Specific components of the blood include red blood cells, white blood cells, platelets, and the plasma, which contains coagulation factors and serum. Blood is important for regulation of the body’s pH, temperature, osmotic pressure, the circulation of nutrients and removal of waste, the distribution of hormones from endocrine glands, and the elimination of excess heat; it also contains components for blood clotting. Red blood cells are specialized cells that contain hemoglobin and circulate through the body delivering oxygen to cells. White blood cells are involved in the immune response to identify and target invading bacteria, viruses, and other foreign organisms; they also recycle waste components, such as old red blood cells. Platelets and blood clotting factors cause the change of the soluble protein fibrinogen to the insoluble protein fibrin at a wound site forming a plug. Plasma consists of 90 percent water along with various substances, such as coagulation factors and antibodies. The serum is the plasma component of the blood without the coagulation factors.
Glossary
plasma
liquid component of blood that is left after the cells are removed
platelet
(also, thrombocyte) small cellular fragment that collects at wounds, cross-reacts with clotting factors, and forms a plug to prevent blood loss
red blood cell
small (7–8 μm) biconcave cell without mitochondria (and in mammals without nuclei) that is packed with hemoglobin, giving the cell its red color; transports oxygen through the body
serum
plasma without the coagulation factors
white blood cell
large (30 μm) cell with nuclei of which there are many types with different roles including the protection of the body from viruses and bacteria, and cleaning up dead cells and other waste | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/48%3A_The_Circulatory_System/48.02%3A_Components_of_Vertebrate_Blood.txt |
Skills to Develop
• Describe the structure of the heart and explain how cardiac muscle is different from other muscles
• Describe the cardiac cycle
• Explain the structure of arteries, veins, and capillaries, and how blood flows through the body
The heart is a complex muscle that pumps blood through the three divisions of the circulatory system: the coronary (vessels that serve the heart), pulmonary (heart and lungs), and systemic (systems of the body), as shown in Figure \(1\). Coronary circulation intrinsic to the heart takes blood directly from the main artery (aorta) coming from the heart. For pulmonary and systemic circulation, the heart has to pump blood to the lungs or the rest of the body, respectively. In vertebrates, the lungs are relatively close to the heart in the thoracic cavity. The shorter distance to pump means that the muscle wall on the right side of the heart is not as thick as the left side which must have enough pressure to pump blood all the way to your big toe.
Exercise
Which of the following statements about the circulatory system is false?
1. Blood in the pulmonary vein is deoxygenated.
2. Blood in the inferior vena cava is deoxygenated.
3. Blood in the pulmonary artery is deoxygenated.
4. Blood in the aorta is oxygenated.
Answer
C
Structure of the Heart
The heart muscle is asymmetrical as a result of the distance blood must travel in the pulmonary and systemic circuits. Since the right side of the heart sends blood to the pulmonary circuit it is smaller than the left side which must send blood out to the whole body in the systemic circuit, as shown in Figure \(2\). In humans, the heart is about the size of a clenched fist; it is divided into four chambers: two atria and two ventricles. There is one atrium and one ventricle on the right side and one atrium and one ventricle on the left side. The atria are the chambers that receive blood, and the ventricles are the chambers that pump blood. The right atrium receives deoxygenated blood from the superior vena cava, which drains blood from the jugular vein that comes from the brain and from the veins that come from the arms, as well as from the inferior vena cava which drains blood from the veins that come from the lower organs and the legs. In addition, the right atrium receives blood from the coronary sinus which drains deoxygenated blood from the heart itself. This deoxygenated blood then passes to the right ventricle through the atrioventricular valve or the tricuspid valve, a flap of connective tissue that opens in only one direction to prevent the backflow of blood.
The valve separating the chambers on the left side of the heart valve is called the biscuspid or mitral valve. After it is filled, the right ventricle pumps the blood through the pulmonary arteries, by-passing the semilunar valve (or pulmonic valve) to the lungs for re-oxygenation. After blood passes through the pulmonary arteries, the right semilunar valves close preventing the blood from flowing backwards into the right ventricle. The left atrium then receives the oxygen-rich blood from the lungs via the pulmonary veins. This blood passes through the bicuspid valve or mitral valve (the atrioventricular valve on the left side of the heart) to the left ventricle where the blood is pumped out through aorta, the major artery of the body, taking oxygenated blood to the organs and muscles of the body. Once blood is pumped out of the left ventricle and into the aorta, the aortic semilunar valve (or aortic valve) closes preventing blood from flowing backward into the left ventricle. This pattern of pumping is referred to as double circulation and is found in all mammals.
Exercise
Which of the following statements about the heart is false?
1. The mitral valve separates the left ventricle from the left atrium.
2. Blood travels through the bicuspid valve to the left atrium.
3. Both the aortic and the pulmonary valves are semilunar valves.
4. The mitral valve is an atrioventricular valve.
Answer
B
The heart is composed of three layers; the epicardium, the myocardium, and the endocardium, illustrated in Figure \(2\). The inner wall of the heart has a lining called the endocardium. The myocardium consists of the heart muscle cells that make up the middle layer and the bulk of the heart wall. The outer layer of cells is called the epicardium, of which the second layer is a membranous layered structure called the pericardium that surrounds and protects the heart; it allows enough room for vigorous pumping but also keeps the heart in place to reduce friction between the heart and other structures.
The heart has its own blood vessels that supply the heart muscle with blood. The coronary arteries branch from the aorta and surround the outer surface of the heart like a crown. They diverge into capillaries where the heart muscle is supplied with oxygen before converging again into the coronary veins to take the deoxygenated blood back to the right atrium where the blood will be re-oxygenated through the pulmonary circuit. The heart muscle will die without a steady supply of blood. Atherosclerosis is the blockage of an artery by the buildup of fatty plaques. Because of the size (narrow) of the coronary arteries and their function in serving the heart itself, atherosclerosis can be deadly in these arteries. The slowdown of blood flow and subsequent oxygen deprivation that results from atherosclerosis causes severe pain, known as angina, and complete blockage of the arteries will cause myocardial infarction: the death of cardiac muscle tissue, commonly known as a heart attack.
The Cardiac Cycle
The main purpose of the heart is to pump blood through the body; it does so in a repeating sequence called the cardiac cycle. The cardiac cycle is the coordination of the filling and emptying of the heart of blood by electrical signals that cause the heart muscles to contract and relax. The human heart beats over 100,000 times per day. In each cardiac cycle, the heart contracts (systole), pushing out the blood and pumping it through the body; this is followed by a relaxation phase (diastole), where the heart fills with blood, as illustrated in Figure \(3\). The atria contract at the same time, forcing blood through the atrioventricular valves into the ventricles. Closing of the atrioventricular valves produces a monosyllabic “lup” sound. Following a brief delay, the ventricles contract at the same time forcing blood through the semilunar valves into the aorta and the artery transporting blood to the lungs (via the pulmonary artery). Closing of the semilunar valves produces a monosyllabic “dup” sound.
The pumping of the heart is a function of the cardiac muscle cells, or cardiomyocytes, that make up the heart muscle. Cardiomyocytes, shown in Figure \(4\), are distinctive muscle cells that are striated like skeletal muscle but pump rhythmically and involuntarily like smooth muscle; they are connected by intercalated disks exclusive to cardiac muscle. They are self-stimulated for a period of time and isolated cardiomyocytes will beat if given the correct balance of nutrients and electrolytes.
The autonomous beating of cardiac muscle cells is regulated by the heart’s internal pacemaker that uses electrical signals to time the beating of the heart. The electrical signals and mechanical actions, illustrated in Figure \(5\), are intimately intertwined. The internal pacemaker starts at the sinoatrial (SA) node, which is located near the wall of the right atrium. Electrical charges spontaneously pulse from the SA node causing the two atria to contract in unison. The pulse reaches a second node, called the atrioventricular (AV) node, between the right atrium and right ventricle where it pauses for approximately 0.1 second before spreading to the walls of the ventricles. From the AV node, the electrical impulse enters the bundle of His, then to the left and right bundle branches extending through the interventricular septum. Finally, the Purkinje fibers conduct the impulse from the apex of the heart up the ventricular myocardium, and then the ventricles contract. This pause allows the atria to empty completely into the ventricles before the ventricles pump out the blood. The electrical impulses in the heart produce electrical currents that flow through the body and can be measured on the skin using electrodes. This information can be observed as an electrocardiogram (ECG)—a recording of the electrical impulses of the cardiac muscle.
Link to Learning
Visit this site and select the dropdown “Your Heart’s Electrical System” to see the heart’s “pacemaker” in action.
Arteries, Veins, and Capillaries
The blood from the heart is carried through the body by a complex network of blood vessels (Figure \(6\)). Arteries take blood away from the heart. The main artery is the aorta that branches into major arteries that take blood to different limbs and organs. These major arteries include the carotid artery that takes blood to the brain, the brachial arteries that take blood to the arms, and the thoracic artery that takes blood to the thorax and then into the hepatic, renal, and gastric arteries for the liver, kidney, and stomach, respectively. The iliac artery takes blood to the lower limbs. The major arteries diverge into minor arteries, and then smaller vessels called arterioles, to reach more deeply into the muscles and organs of the body.
Arterioles diverge into capillary beds. Capillary beds contain a large number (10 to 100) of capillaries that branch among the cells and tissues of the body. Capillaries are narrow-diameter tubes that can fit red blood cells through in single file and are the sites for the exchange of nutrients, waste, and oxygen with tissues at the cellular level. Fluid also crosses into the interstitial space from the capillaries. The capillaries converge again into venules that connect to minor veins that finally connect to major veins that take blood high in carbon dioxide back to the heart. Veins are blood vessels that bring blood back to the heart. The major veins drain blood from the same organs and limbs that the major arteries supply. Fluid is also brought back to the heart via the lymphatic system.
The structure of the different types of blood vessels reflects their function or layers. There are three distinct layers, or tunics, that form the walls of blood vessels (Figure \(7\)). The first tunic is a smooth, inner lining of endothelial cells that are in contact with the red blood cells. The endothelial tunic is continuous with the endocardium of the heart. In capillaries, this single layer of cells is the location of diffusion of oxygen and carbon dioxide between the endothelial cells and red blood cells, as well as the exchange site via endocytosis and exocytosis. The movement of materials at the site of capillaries is regulated by vasoconstriction, narrowing of the blood vessels, and vasodilation, widening of the blood vessels; this is important in the overall regulation of blood pressure.
Veins and arteries both have two further tunics that surround the endothelium: the middle tunic is composed of smooth muscle and the outermost layer is connective tissue (collagen and elastic fibers). The elastic connective tissue stretches and supports the blood vessels, and the smooth muscle layer helps regulate blood flow by altering vascular resistance through vasoconstriction and vasodilation. The arteries have thicker smooth muscle and connective tissue than the veins to accommodate the higher pressure and speed of freshly pumped blood. The veins are thinner walled as the pressure and rate of flow are much lower. In addition, veins are structurally different than arteries in that veins have valves to prevent the backflow of blood. Because veins have to work against gravity to get blood back to the heart, contraction of skeletal muscle assists with the flow of blood back to the heart.
Summary
The heart muscle pumps blood through three divisions of the circulatory system: coronary, pulmonary, and systemic. There is one atrium and one ventricle on the right side and one atrium and one ventricle on the left side. The pumping of the heart is a function of cardiomyocytes, distinctive muscle cells that are striated like skeletal muscle but pump rhythmically and involuntarily like smooth muscle. The internal pacemaker starts at the sinoatrial node, which is located near the wall of the right atrium. Electrical charges pulse from the SA node causing the two atria to contract in unison; then the pulse reaches the atrioventricular node between the right atrium and right ventricle. A pause in the electric signal allows the atria to empty completely into the ventricles before the ventricles pump out the blood. The blood from the heart is carried through the body by a complex network of blood vessels; arteries take blood away from the heart, and veins bring blood back to the heart.
Glossary
angina
pain caused by partial blockage of the coronary arteries by the buildup of plaque and lack of oxygen to the heart muscle
aorta
major artery of the body that takes blood away from the heart
arteriole
small vessel that connects an artery to a capillary bed
artery
blood vessel that takes blood away from the heart
atherosclerosis
buildup of fatty plaques in the coronary arteries in the heart
atrioventricular valve
one-way membranous flap of connective tissue between the atrium and the ventricle in the right side of the heart; also known as tricuspid valve
bicuspid valve
(also, mitral valve; left atrioventricular valve) one-way membranous flap between the atrium and the ventricle in the left side of the heart
capillary
smallest blood vessel that allows the passage of individual blood cells and the site of diffusion of oxygen and nutrient exchange
capillary bed
large number of capillaries that converge to take blood to a particular organ or tissue
cardiac cycle
filling and emptying the heart of blood by electrical signals that cause the heart muscles to contract and relax
cardiomyocyte
specialized heart muscle cell that is striated but contracts involuntarily like smooth muscle
coronary artery
vessel that supplies the heart tissue with blood
coronary vein
vessel that takes blood away from the heart tissue back to the chambers in the heart
diastole
relaxation phase of the cardiac cycle when the heart is relaxed and the ventricles are filling with blood
electrocardiogram (ECG)
recording of the electrical impulses of the cardiac muscle
endocardium
innermost layer of tissue in the heart
epicardium
outermost tissue layer of the heart
inferior vena cava
drains blood from the veins that come from the lower organs and the legs
myocardial infarction
(also, heart attack) complete blockage of the coronary arteries and death of the cardiac muscle tissue
myocardium
heart muscle cells that make up the middle layer and the bulk of the heart wall
pericardium
membrane layer protecting the heart; also part of the epicardium
semilunar valve
membranous flap of connective tissue between the aorta and a ventricle of the heart (the aortic or pulmonary semilunar valves)
sinoatrial (SA) node
the heart’s internal pacemaker; located near the wall of the right atrium
superior vena cava
drains blood from the jugular vein that comes from the brain and from the veins that come from the arms
systole
contraction phase of cardiac cycle when the ventricles are pumping blood into the arteries
tricuspid valve
one-way membranous flap of connective tissue between the atrium and the ventricle in the right side of the heart; also known as atrioventricular valve
vasoconstriction
narrowing of a blood vessel
vasodilation
widening of a blood vessel
vein
blood vessel that brings blood back to the heart
vena cava
major vein of the body returning blood from the upper and lower parts of the body; see the superior vena cava and inferior vena cava
venule
blood vessel that connects a capillary bed to a vein | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/48%3A_The_Circulatory_System/48.03%3A_Vertebrate_Circulatory_System.txt |
Skills to Develop
• Describe the system of blood flow through the body
• Describe how blood pressure is regulated
Blood pressure (BP) is the pressure exerted by blood on the walls of a blood vessel that helps to push blood through the body. Systolic blood pressure measures the amount of pressure that blood exerts on vessels while the heart is beating. The optimal systolic blood pressure is 120 mmHg. Diastolic blood pressure measures the pressure in the vessels between heartbeats. The optimal diastolic blood pressure is 80 mmHg. Many factors can affect blood pressure, such as hormones, stress, exercise, eating, sitting, and standing. Blood flow through the body is regulated by the size of blood vessels, by the action of smooth muscle, by one-way valves, and by the fluid pressure of the blood itself.
How Blood Flows Through the Body
Blood is pushed through the body by the action of the pumping heart. With each rhythmic pump, blood is pushed under high pressure and velocity away from the heart, initially along the main artery, the aorta. In the aorta, the blood travels at 30 cm/sec. As blood moves into the arteries, arterioles, and ultimately to the capillary beds, the rate of movement slows dramatically to about 0.026 cm/sec, one-thousand times slower than the rate of movement in the aorta. While the diameter of each individual arteriole and capillary is far narrower than the diameter of the aorta, and according to the law of continuity, fluid should travel faster through a narrower diameter tube, the rate is actually slower due to the overall diameter of all the combined capillaries being far greater than the diameter of the individual aorta.
The slow rate of travel through the capillary beds, which reach almost every cell in the body, assists with gas and nutrient exchange and also promotes the diffusion of fluid into the interstitial space. After the blood has passed through the capillary beds to the venules, veins, and finally to the main venae cavae, the rate of flow increases again but is still much slower than the initial rate in the aorta. Blood primarily moves in the veins by the rhythmic movement of smooth muscle in the vessel wall and by the action of the skeletal muscle as the body moves. Because most veins must move blood against the pull of gravity, blood is prevented from flowing backward in the veins by one-way valves. Because skeletal muscle contraction aids in venous blood flow, it is important to get up and move frequently after long periods of sitting so that blood will not pool in the extremities.
Blood flow through the capillary beds is regulated depending on the body’s needs and is directed by nerve and hormone signals. For example, after a large meal, most of the blood is diverted to the stomach by vasodilation of vessels of the digestive system and vasoconstriction of other vessels. During exercise, blood is diverted to the skeletal muscles through vasodilation while blood to the digestive system would be lessened through vasoconstriction. The blood entering some capillary beds is controlled by small muscles, called precapillary sphincters, illustrated in Figure \(1\). If the sphincters are open, the blood will flow into the associated branches of the capillary blood. If all of the sphincters are closed, then the blood will flow directly from the arteriole to the venule through the thoroughfare channel (see Figure \(1\)). These muscles allow the body to precisely control when capillary beds receive blood flow. At any given moment only about 5-10% of our capillary beds actually have blood flowing through them.
Exercise
Varicose veins are veins that become enlarged because the valves no longer close properly, allowing blood to flow backward. Varicose veins are often most prominent on the legs. Why do you think this is the case?
Answer
Blood in the legs is farthest away from the heart and has to flow up to reach it.
Link to Learning
The circulatory system consisting of the heart, arteries, capillaries, and veins, is the pumping mechanism that transports blood throughout the body. Watch this video to see the circulatory system’s blood flow.
Proteins and other large solutes cannot leave the capillaries. The loss of the watery plasma creates a hyperosmotic solution within the capillaries, especially near the venules. This causes about 85% of the plasma that leaves the capillaries to eventually diffuses back into the capillaries near the venules. The remaining 15% of blood plasma drains out from the interstitial fluid into nearby lymphatic vessels (Figure \(2\)). The fluid in the lymph is similar in composition to the interstitial fluid. The lymph fluid passes through lymph nodes before it returns to the heart via the vena cava. Lymph nodes are specialized organs that filter the lymph by percolation through a maze of connective tissue filled with white blood cells. The white blood cells remove infectious agents, such as bacteria and viruses, to clean the lymph before it returns to the bloodstream. After it is cleaned, the lymph returns to the heart by the action of smooth muscle pumping, skeletal muscle action, and one-way valves joining the returning blood near the junction of the venae cavae entering the right atrium of the heart.
Evolution Connection: Vertebrate Diversity in Blood Circulation
Blood circulation has evolved differently in vertebrates and may show variation in different animals for the required amount of pressure, organ and vessel location, and organ size. Animals with longs necks and those that live in cold environments have distinct blood pressure adaptations.
Long necked animals, such as giraffes, need to pump blood upward from the heart against gravity. The blood pressure required from the pumping of the left ventricle would be equivalent to 250 mm Hg (mm Hg = millimeters of mercury, a unit of pressure) to reach the height of a giraffe’s head, which is 2.5 meters higher than the heart. However, if checks and balances were not in place, this blood pressure would damage the giraffe’s brain, particularly if it was bending down to drink. These checks and balances include valves and feedback mechanisms that reduce the rate of cardiac output. Long-necked dinosaurs such as the sauropods had to pump blood even higher, up to ten meters above the heart. This would have required a blood pressure of more than 600 mm Hg, which could only have been achieved by an enormous heart. Evidence for such an enormous heart does not exist and mechanisms to reduce the blood pressure required include the slowing of metabolism as these animals grew larger. It is likely that they did not routinely feed on tree tops but grazed on the ground.
Living in cold water, whales need to maintain the temperature in their blood. This is achieved by the veins and arteries being close together so that heat exchange can occur. This mechanism is called a countercurrent heat exchanger. The blood vessels and the whole body are also protected by thick layers of blubber to prevent heat loss. In land animals that live in cold environments, thick fur and hibernation are used to retain heat and slow metabolism.
Blood Pressure
The pressure of the blood flow in the body is produced by the hydrostatic pressure of the fluid (blood) against the walls of the blood vessels. Fluid will move from areas of high to low hydrostatic pressures. In the arteries, the hydrostatic pressure near the heart is very high and blood flows to the arterioles where the rate of flow is slowed by the narrow openings of the arterioles. During systole, when new blood is entering the arteries, the artery walls stretch to accommodate the increase of pressure of the extra blood; during diastole, the walls return to normal because of their elastic properties. The blood pressure of the systole phase and the diastole phase, graphed in Figure \(3\), gives the two pressure readings for blood pressure. For example, 120/80 indicates a reading of 120 mm Hg during the systole and 80 mm Hg during diastole. Throughout the cardiac cycle, the blood continues to empty into the arterioles at a relatively even rate. This resistance to blood flow is called peripheral resistance.
Blood Pressure Regulation
Cardiac output is the volume of blood pumped by the heart in one minute. It is calculated by multiplying the number of heart contractions that occur per minute (heart rate) times the stroke volume (the volume of blood pumped into the aorta per contraction of the left ventricle). Therefore, cardiac output can be increased by increasing heart rate, as when exercising. However, cardiac output can also be increased by increasing stroke volume, such as if the heart contracts with greater strength. Stroke volume can also be increased by speeding blood circulation through the body so that more blood enters the heart between contractions. During heavy exertion, the blood vessels relax and increase in diameter, offsetting the increased heart rate and ensuring adequate oxygenated blood gets to the muscles. Stress triggers a decrease in the diameter of the blood vessels, consequently increasing blood pressure. These changes can also be caused by nerve signals or hormones, and even standing up or lying down can have a great effect on blood pressure.
Summary
Blood primarily moves through the body by the rhythmic movement of smooth muscle in the vessel wall and by the action of the skeletal muscle as the body moves. Blood is prevented from flowing backward in the veins by one-way valves. Blood flow through the capillary beds is controlled by precapillary sphincters to increase and decrease flow depending on the body’s needs and is directed by nerve and hormone signals. Lymph vessels take fluid that has leaked out of the blood to the lymph nodes where it is cleaned before returning to the heart. During systole, blood enters the arteries, and the artery walls stretch to accommodate the extra blood. During diastole, the artery walls return to normal. The blood pressure of the systole phase and the diastole phase gives the two pressure readings for blood pressure.
Glossary
blood pressure (BP)
pressure of blood in the arteries that helps to push blood through the body
cardiac output
the volume of blood pumped by the heart in one minute as a product of heart rate multiplied by stroke volume
lymph node
specialized organ that contains a large number of macrophages that clean the lymph before the fluid is returned to the heart
peripheral resistance
resistance of the artery and blood vessel walls to the pressure placed on them by the force of the heart pumping
precapillary sphincter
small muscle that controls blood circulation in the capillary beds
stroke volume
the volume of blood pumped into the aorta per contraction of the left ventricle | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/48%3A_The_Circulatory_System/48.04%3A_Cardiac_Cycle_Electrical_Conduction_ECG_and_Cardiac_Output.txt |
• 49.1: Osmolarity and Osmotic Balance
Osmosis is the diffusion of water across a membrane in response to osmotic pressure caused by an imbalance of molecules on either side of the membrane. Osmoregulation is the process of maintenance of salt and water balance (osmotic balance) across membranes within the body’s fluids, which are composed of water, plus electrolytes and non-electrolytes.
• 49.2: Nitrogenous Wastes- Ammonia, Urea, and Uric Acid
Of the four major macromolecules in biological systems, both proteins and nucleic acids contain nitrogen. During the catabolism, or breakdown, of nitrogen-containing macromolecules, carbon, hydrogen, and oxygen are extracted and stored in the form of carbohydrates and fats. Excess nitrogen is excreted from the body. Nitrogenous wastes tend to form toxic ammonia, which raises the pH of body fluids.
• 49.3: Osmoregulatory Organs
Although the kidneys are the major osmoregulatory organ, the skin and lungs also play a role in the process. Water and electrolytes are lost through sweat glands in the skin, which helps moisturize and cool the skin surface, while the lungs expel a small amount of water in the form of mucous secretions and via evaporation of water vapor.
• 49.4: Evolution of the Vertebrate Kidney
• 49.5: The Mammalian Kidney
Although the kidneys are the major osmoregulatory organ, the skin and lungs also play a role in the process. Water and electrolytes are lost through sweat glands in the skin, which helps moisturize and cool the skin surface, while the lungs expel a small amount of water in the form of mucous secretions and via evaporation of water vapor.
• 49.6: Hormonal Control of Osmoregulatory Functions
While the kidneys operate to maintain osmotic balance and blood pressure in the body, they also act in concert with hormones. Hormones are small molecules that act as messengers within the body. Hormones are typically secreted from one cell and travel in the bloodstream to affect a target cell in another portion of the body. Different regions of the nephron bear specialized cells that have receptors to respond to chemical messengers and hormones.
49: Osmotic Regulation and the Urinary System
Skills to Develop
• Define osmosis and explain its role within molecules
• Explain why osmoregulation and osmotic balance are important body functions
• Describe active transport mechanisms
• Explain osmolarity and the way in which it is measured
• Describe osmoregulators or osmoconformers and how these tools allow animals to adapt to different environments
Osmosis is the diffusion of water across a membrane in response to osmotic pressure caused by an imbalance of molecules on either side of the membrane. Osmoregulation is the process of maintenance of salt and water balance (osmotic balance) across membranes within the body’s fluids, which are composed of water, plus electrolytes and non-electrolytes. An electrolyte is a solute that dissociates into ions when dissolved in water. A non-electrolyte, in contrast, doesn’t dissociate into ions during water dissolution. Both electrolytes and non-electrolytes contribute to the osmotic balance. The body’s fluids include blood plasma, the cytosol within cells, and interstitial fluid, the fluid that exists in the spaces between cells and tissues of the body. The membranes of the body (such as the pleural, serous, and cell membranes) are semi-permeable membranes. Semi-permeable membranes are permeable (or permissive) to certain types of solutes and water. Solutions on two sides of a semi-permeable membrane tend to equalize in solute concentration by movement of solutes and/or water across the membrane. As seen in Figure \(1\), a cell placed in water tends to swell due to gain of water from the hypotonic or “low salt” environment. A cell placed in a solution with higher salt concentration, on the other hand, tends to make the membrane shrivel up due to loss of water into the hypertonic or “high salt” environment. Isotonic cells have an equal concentration of solutes inside and outside the cell; this equalizes the osmotic pressure on either side of the cell membrane which is a semi-permeable membrane.
The body does not exist in isolation. There is a constant input of water and electrolytes into the system. While osmoregulation is achieved across membranes within the body, excess electrolytes and wastes are transported to the kidneys and excreted, helping to maintain osmotic balance.
Need for Osmoregulation
Biological systems constantly interact and exchange water and nutrients with the environment by way of consumption of food and water and through excretion in the form of sweat, urine, and feces. Without a mechanism to regulate osmotic pressure, or when a disease damages this mechanism, there is a tendency to accumulate toxic waste and water, which can have dire consequences.
Mammalian systems have evolved to regulate not only the overall osmotic pressure across membranes, but also specific concentrations of important electrolytes in the three major fluid compartments: blood plasma, extracellular fluid, and intracellular fluid. Since osmotic pressure is regulated by the movement of water across membranes, the volume of the fluid compartments can also change temporarily. Because blood plasma is one of the fluid components, osmotic pressures have a direct bearing on blood pressure.
Transport of Electrolytes across Cell Membranes
Electrolytes, such as sodium chloride, ionize in water, meaning that they dissociate into their component ions. In water, sodium chloride (NaCl), dissociates into the sodium ion (Na+) and the chloride ion (Cl). The most important ions, whose concentrations are very closely regulated in body fluids, are the cations sodium (Na+), potassium (K+), calcium (Ca+2), magnesium (Mg+2), and the anions chloride (Cl-), carbonate (CO3-2), bicarbonate (HCO3-), and phosphate(PO3-). Electrolytes are lost from the body during urination and perspiration. For this reason, athletes are encouraged to replace electrolytes and fluids during periods of increased activity and perspiration.
Osmotic pressure is influenced by the concentration of solutes in a solution. It is directly proportional to the number of solute atoms or molecules and not dependent on the size of the solute molecules. Because electrolytes dissociate into their component ions, they, in essence, add more solute particles into the solution and have a greater effect on osmotic pressure, per mass than compounds that do not dissociate in water, such as glucose.
Water can pass through membranes by passive diffusion. If electrolyte ions could passively diffuse across membranes, it would be impossible to maintain specific concentrations of ions in each fluid compartment therefore they require special mechanisms to cross the semi-permeable membranes in the body. This movement can be accomplished by facilitated diffusion and active transport. Facilitated diffusion requires protein-based channels for moving the solute. Active transport requires energy in the form of ATP conversion, carrier proteins, or pumps in order to move ions against the concentration gradient.
Concept of Osmolality and Milliequivalent
In order to calculate osmotic pressure, it is necessary to understand how solute concentrations are measured. The unit for measuring solutes is the mole. One mole is defined as the gram molecular weight of the solute. For example, the molecular weight of sodium chloride is 58.44. Thus, one mole of sodium chloride weighs 58.44 grams. The molarity of a solution is the number of moles of solute per liter of solution. The molality of a solution is the number of moles of solute per kilogram of solvent. If the solvent is water, one kilogram of water is equal to one liter of water. While molarity and molality are used to express the concentration of solutions, electrolyte concentrations are usually expressed in terms of milliequivalents per liter (mEq/L): the mEq/L is equal to the ion concentration (in millimoles) multiplied by the number of electrical charges on the ion. The unit of milliequivalent takes into consideration the ions present in the solution (since electrolytes form ions in aqueous solutions) and the charge on the ions.
Thus, for ions that have a charge of one, one milliequivalent is equal to one millimole. For ions that have a charge of two (like calcium), one milliequivalent is equal to 0.5 millimoles. Another unit for the expression of electrolyte concentration is the milliosmole (mOsm), which is the number of milliequivalents of solute per kilogram of solvent. Body fluids are usually maintained within the range of 280 to 300 mOsm.
Osmoregulators and Osmoconformers
Persons lost at sea without any fresh water to drink are at risk of severe dehydration because the human body cannot adapt to drinking seawater, which is hypertonic in comparison to body fluids. Organisms such as goldfish that can tolerate only a relatively narrow range of salinity are referred to as stenohaline. About 90 percent of all bony fish are restricted to either freshwater or seawater. They are incapable of osmotic regulation in the opposite environment. It is possible, however, for a few fishes like salmon to spend part of their life in fresh water and part in sea water. Organisms like the salmon and molly that can tolerate a relatively wide range of salinity are referred to as euryhaline organisms. This is possible because some fish have evolved osmoregulatory mechanisms to survive in all kinds of aquatic environments. When they live in fresh water, their bodies tend to take up water because the environment is relatively hypotonic, as illustrated in Figure \(2\)a. In such hypotonic environments, these fish do not drink much water. Instead, they pass a lot of very dilute urine, and they achieve electrolyte balance by active transport of salts through the gills. When they move to a hypertonic marine environment, these fish start drinking sea water; they excrete the excess salts through their gills and their urine, as illustrated in Figure \(2\)b. Most marine invertebrates, on the other hand, may be isotonic with sea water (osmoconformers). Their body fluid concentrations conform to changes in seawater concentration. Cartilaginous fishes’ salt composition of the blood is similar to bony fishes; however, the blood of sharks contains the organic compounds urea and trimethylamine oxide (TMAO). This does not mean that their electrolyte composition is similar to that of sea water. They achieve isotonicity with the sea by storing large concentrations of urea. These animals that secrete urea are called ureotelic animals. TMAO stabilizes proteins in the presence of high urea levels, preventing the disruption of peptide bonds that would occur in other animals exposed to similar levels of urea. Sharks are cartilaginous fish with a rectal gland to secrete salt and assist in osmoregulation.
Career Connection: Dialysis Technician
Dialysis is a medical process of removing wastes and excess water from the blood by diffusion and ultrafiltration. When kidney function fails, dialysis must be done to artificially rid the body of wastes. This is a vital process to keep patients alive. In some cases, the patients undergo artificial dialysis until they are eligible for a kidney transplant. In others who are not candidates for kidney transplants, dialysis is a life-long necessity.
Dialysis technicians typically work in hospitals and clinics. While some roles in this field include equipment development and maintenance, most dialysis technicians work in direct patient care. Their on-the-job duties, which typically occur under the direct supervision of a registered nurse, focus on providing dialysis treatments. This can include reviewing patient history and current condition, assessing and responding to patient needs before and during treatment, and monitoring the dialysis process. Treatment may include taking and reporting a patient’s vital signs and preparing solutions and equipment to ensure accurate and sterile procedures.
Summary
Solute concentrations across a semi-permeable membranes influence the movement of water and solutes across the membrane. It is the number of solute molecules and not the molecular size that is important in osmosis. Osmoregulation and osmotic balance are important bodily functions, resulting in water and salt balance. Not all solutes can pass through a semi-permeable membrane. Osmosis is the movement of water across the membrane. Osmosis occurs to equalize the number of solute molecules across a semi-permeable membrane by the movement of water to the side of higher solute concentration. Facilitated diffusion utilizes protein channels to move solute molecules from areas of higher to lower concentration while active transport mechanisms are required to move solutes against concentration gradients. Osmolarity is measured in units of milliequivalents or milliosmoles, both of which take into consideration the number of solute particles and the charge on them. Fish that live in fresh water or saltwater adapt by being osmoregulators or osmoconformers.
Glossary
electrolyte
solute that breaks down into ions when dissolved in water
molality
number of moles of solute per kilogram of solvent
molarity
number of moles of solute per liter of solution
mole
gram equivalent of the molecular weight of a substance
non-electrolyte
solute that does not break down into ions when dissolved in water
osmoconformer
organism that changes its tonicity based on its environment
osmoregulation
mechanism by which water and solute concentrations are maintained at desired levels
osmoregulator
organism that maintains its tonicity irrespective of its environment
osmotic balance
balance of the amount of water and salt input and output to and from a biological system without disturbing the desired osmotic pressure and solute concentration in every compartment
osmotic pressure
pressure exerted on a membrane to equalize solute concentration on either side
semi-permeable membrane
membrane that allows only certain solutes to pass through | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/49%3A_Osmotic_Regulation_and_the_Urinary_System/49.01%3A_Osmolarity_and_Osmotic_Balance.txt |
Skills to Develop
• Compare and contrast the way in which aquatic animals and terrestrial animals can eliminate toxic ammonia from their systems
• Compare the major byproduct of ammonia metabolism in vertebrate animals to that of birds, insects, and reptiles
Of the four major macromolecules in biological systems, both proteins and nucleic acids contain nitrogen. During the catabolism, or breakdown, of nitrogen-containing macromolecules, carbon, hydrogen, and oxygen are extracted and stored in the form of carbohydrates and fats. Excess nitrogen is excreted from the body. Nitrogenous wastes tend to form toxic ammonia, which raises the pH of body fluids. The formation of ammonia itself requires energy in the form of ATP and large quantities of water to dilute it out of a biological system. Animals that live in aquatic environments tend to release ammonia into the water. Animals that excrete ammonia are said to be ammonotelic. Terrestrial organisms have evolved other mechanisms to excrete nitrogenous wastes. The animals must detoxify ammonia by converting it into a relatively nontoxic form such as urea or uric acid. Mammals, including humans, produce urea, whereas reptiles and many terrestrial invertebrates produce uric acid. Animals that secrete urea as the primary nitrogenous waste material are called ureotelic animals.
Nitrogenous Waste in Terrestrial Animals: The Urea Cycle
The urea cycle is the primary mechanism by which mammals convert ammonia to urea. Urea is made in the liver and excreted in urine. The overall chemical reaction by which ammonia is converted to urea is 2 NH3 (ammonia) + CO2 + 3 ATP + H2O → H2N-CO-NH2 (urea) + 2 ADP + 4 Pi + AMP.
The urea cycle utilizes five intermediate steps, catalyzed by five different enzymes, to convert ammonia to urea, as shown in Figure \(1\). The amino acid L-ornithine gets converted into different intermediates before being regenerated at the end of the urea cycle. Hence, the urea cycle is also referred to as the ornithine cycle. The enzyme ornithine transcarbamylase catalyzes a key step in the urea cycle and its deficiency can lead to accumulation of toxic levels of ammonia in the body. The first two reactions occur in the mitochondria and the last three reactions occur in the cytosol. Urea concentration in the blood, called blood urea nitrogen or BUN, is used as an indicator of kidney function.
Evolution Connection: Excretion of Nitrogenous Waste
The theory of evolution proposes that life started in an aquatic environment. It is not surprising to see that biochemical pathways like the urea cycle evolved to adapt to a changing environment when terrestrial life forms evolved. Arid conditions probably led to the evolution of the uric acid pathway as a means of conserving water.
Nitrogenous Waste in Birds and Reptiles: Uric Acid
Birds, reptiles, and most terrestrial arthropods convert toxic ammonia to uric acid or the closely related compound guanine (guano) instead of urea. Mammals also form some uric acid during breakdown of nucleic acids. Uric acid is a compound similar to purines found in nucleic acids. It is water insoluble and tends to form a white paste or powder; it is excreted by birds, insects, and reptiles. Conversion of ammonia to uric acid requires more energy and is much more complex than conversion of ammonia to urea Figure \(2\).
Everyday Connection: Gout
Mammals use uric acid crystals as an antioxidant in their cells. However, too much uric acid tends to form kidney stones and may also cause a painful condition called gout, where uric acid crystals accumulate in the joints, as illustrated in Figure \(3\). Food choices that reduce the amount of nitrogenous bases in the diet help reduce the risk of gout. For example, tea, coffee, and chocolate have purine-like compounds, called xanthines, and should be avoided by people with gout and kidney stones.
Summary
Ammonia is the waste produced by metabolism of nitrogen-containing compounds like proteins and nucleic acids. While aquatic animals can easily excrete ammonia into their watery surroundings, terrestrial animals have evolved special mechanisms to eliminate the toxic ammonia from their systems. Urea is the major byproduct of ammonia metabolism in vertebrate animals. Uric acid is the major byproduct of ammonia metabolism in birds, terrestrial arthropods, and reptiles.
Glossary
ammonia
compound made of one nitrogen atom and three hydrogen atoms
ammonotelic
describes an animal that excretes ammonia as the primary waste material
antioxidant
agent that prevents cell destruction by reactive oxygen species
blood urea nitrogen (BUN)
estimate of urea in the blood and an indicator of kidney function
urea cycle
pathway by which ammonia is converted to urea
ureotelic
describes animals that secrete urea as the primary nitrogenous waste material
uric acid
byproduct of ammonia metabolism in birds, insects, and reptiles | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/49%3A_Osmotic_Regulation_and_the_Urinary_System/49.02%3A_Nitrogenous_Wastes-_Ammonia_Urea_and_Uric_Acid.txt |
Skills to Develop
• Explain how the kidneys serve as the main osmoregulatory organs in mammalian systems
• Describe the structure of the kidneys and the functions of the parts of the kidney
• Describe how the nephron is the functional unit of the kidney and explain how it actively filters blood and generates urine
• Detail the three steps in the formation of urine: glomerular filtration, tubular reabsorption, and tubular secretion
Although the kidneys are the major osmoregulatory organ, the skin and lungs also play a role in the process. Water and electrolytes are lost through sweat glands in the skin, which helps moisturize and cool the skin surface, while the lungs expel a small amount of water in the form of mucous secretions and via evaporation of water vapor.
Kidneys: The Main Osmoregulatory Organ
The kidneys, illustrated in Figure \(1\), are a pair of bean-shaped structures that are located just below and posterior to the liver in the peritoneal cavity. The adrenal glands sit on top of each kidney and are also called the suprarenal glands. Kidneys filter blood and purify it. All the blood in the human body is filtered many times a day by the kidneys; these organs use up almost 25 percent of the oxygen absorbed through the lungs to perform this function. Oxygen allows the kidney cells to efficiently manufacture chemical energy in the form of ATP through aerobic respiration. The filtrate coming out of the kidneys is called urine.
Kidney Structure
Externally, the kidneys are surrounded by three layers, illustrated in Figure \(2\). The outermost layer is a tough connective tissue layer called the renal fascia. The second layer is called the perirenal fat capsule, which helps anchor the kidneys in place. The third and innermost layer is the renal capsule. Internally, the kidney has three regions—an outer cortex, a medulla in the middle, and the renal pelvis in the region called the hilum of the kidney. The hilum is the concave part of the bean-shape where blood vessels and nerves enter and exit the kidney; it is also the point of exit for the ureters. The renal cortex is granular due to the presence of nephrons—the functional unit of the kidney. The medulla consists of multiple pyramidal tissue masses, called the renal pyramids. In between the pyramids are spaces called renal columns through which the blood vessels pass. The tips of the pyramids, called renal papillae, point toward the renal pelvis. There are, on average, eight renal pyramids in each kidney. The renal pyramids along with the adjoining cortical region are called the lobes of the kidney. The renal pelvis leads to the ureter on the outside of the kidney. On the inside of the kidney, the renal pelvis branches out into two or three extensions called the major calyces, which further branch into the minor calyces. The ureters are urine-bearing tubes that exit the kidney and empty into the urinary bladder.
Art Connection
Which of the following statements about the kidney is false?
1. The renal pelvis drains into the ureter.
2. The renal pyramids are in the medulla.
3. The cortex covers the capsule.
4. Nephrons are in the renal cortex.
Because the kidney filters blood, its network of blood vessels is an important component of its structure and function. The arteries, veins, and nerves that supply the kidney enter and exit at the renal hilum. Renal blood supply starts with the branching of the aorta into the renal arteries (which are each named based on the region of the kidney they pass through) and ends with the exiting of the renal veins to join the inferior vena cava. The renal arteries split into several segmental arteries upon entering the kidneys. Each segmental artery splits further into several interlobar arteries and enters the renal columns, which supply the renal lobes. The interlobar arteries split at the junction of the renal cortex and medulla to form the arcuate arteries. The arcuate “bow shaped” arteries form arcs along the base of the medullary pyramids. Cortical radiate arteries, as the name suggests, radiate out from the arcuate arteries. The cortical radiate arteries branch into numerous afferent arterioles, and then enter the capillaries supplying the nephrons. Veins trace the path of the arteries and have similar names, except there are no segmental veins.
As mentioned previously, the functional unit of the kidney is the nephron, illustrated in Figure \(3\). Each kidney is made up of over one million nephrons that dot the renal cortex, giving it a granular appearance when sectioned sagittally. There are two types of nephrons—cortical nephrons (85 percent), which are deep in the renal cortex, and juxtamedullary nephrons (15 percent), which lie in the renal cortex close to the renal medulla. A nephron consists of three parts—a renal corpuscle, a renal tubule, and the associated capillary network, which originates from the cortical radiate arteries.
Art Connection
Which of the following statements about the nephron is false?
1. The collecting duct empties into the distal convoluted tubule.
2. The Bowman’s capsule surrounds the glomerulus.
3. The loop of Henle is between the proximal and distal convoluted tubules.
4. The loop of Henle empties into the distal convoluted tubule.
Renal Corpuscle
The renal corpuscle, located in the renal cortex, is made up of a network of capillaries known as the glomerulus and the capsule, a cup-shaped chamber that surrounds it, called the glomerular or Bowman's capsule.
Renal Tubule
The renal tubule is a long and convoluted structure that emerges from the glomerulus and can be divided into three parts based on function. The first part is called the proximal convoluted tubule (PCT) due to its proximity to the glomerulus; it stays in the renal cortex. The second part is called the loop of Henle, or nephritic loop, because it forms a loop (with descending and ascending limbs) that goes through the renal medulla. The third part of the renal tubule is called the distal convoluted tubule (DCT) and this part is also restricted to the renal cortex. The DCT, which is the last part of the nephron, connects and empties its contents into collecting ducts that line the medullary pyramids. The collecting ducts amass contents from multiple nephrons and fuse together as they enter the papillae of the renal medulla.
Capillary Network within the Nephron
The capillary network that originates from the renal arteries supplies the nephron with blood that needs to be filtered. The branch that enters the glomerulus is called the afferent arteriole. The branch that exits the glomerulus is called the efferent arteriole. Within the glomerulus, the network of capillaries is called the glomerular capillary bed. Once the efferent arteriole exits the glomerulus, it forms the peritubular capillary network, which surrounds and interacts with parts of the renal tubule. In cortical nephrons, the peritubular capillary network surrounds the PCT and DCT. In juxtamedullary nephrons, the peritubular capillary network forms a network around the loop of Henle and is called the vasa recta.
Kidney Function and Physiology
Kidneys filter blood in a three-step process. First, the nephrons filter blood that runs through the capillary network in the glomerulus. Almost all solutes, except for proteins, are filtered out into the glomerulus by a process called glomerular filtration. Second, the filtrate is collected in the renal tubules. Most of the solutes get reabsorbed in the PCT by a process called tubular reabsorption. In the loop of Henle, the filtrate continues to exchange solutes and water with the renal medulla and the peritubular capillary network. Water is also reabsorbed during this step. Then, additional solutes and wastes are secreted into the kidney tubules during tubular secretion, which is, in essence, the opposite process to tubular reabsorption. The collecting ducts collect filtrate coming from the nephrons and fuse in the medullary papillae. From here, the papillae deliver the filtrate, now called urine, into the minor calyces that eventually connect to the ureters through the renal pelvis. This entire process is illustrated in Figure \(4\).
Glomerular Filtration
Glomerular filtration filters out most of the solutes due to high blood pressure and specialized membranes in the afferent arteriole. The blood pressure in the glomerulus is maintained independent of factors that affect systemic blood pressure. The “leaky” connections between the endothelial cells of the glomerular capillary network allow solutes to pass through easily. All solutes in the glomerular capillaries, except for macromolecules like proteins, pass through by passive diffusion. There is no energy requirement at this stage of the filtration process. Glomerular filtration rate (GFR) is the volume of glomerular filtrate formed per minute by the kidneys. GFR is regulated by multiple mechanisms and is an important indicator of kidney function.
Link to Learning
To learn more about the vascular system of kidneys, click through this review and the steps of blood flow.
Tubular Reabsorption and Secretion
Tubular reabsorption occurs in the PCT part of the renal tubule. Almost all nutrients are reabsorbed, and this occurs either by passive or active transport. Reabsorption of water and some key electrolytes are regulated and can be influenced by hormones. Sodium (Na+) is the most abundant ion and most of it is reabsorbed by active transport and then transported to the peritubular capillaries. Because Na+ is actively transported out of the tubule, water follows it to even out the osmotic pressure. Water is also independently reabsorbed into the peritubular capillaries due to the presence of aquaporins, or water channels, in the PCT. This occurs due to the low blood pressure and high osmotic pressure in the peritubular capillaries. However, every solute has a transport maximum and the excess is not reabsorbed.
In the loop of Henle, the permeability of the membrane changes. The descending limb is permeable to water, not solutes; the opposite is true for the ascending limb. Additionally, the loop of Henle invades the renal medulla, which is naturally high in salt concentration and tends to absorb water from the renal tubule and concentrate the filtrate. The osmotic gradient increases as it moves deeper into the medulla. Because two sides of the loop of Henle perform opposing functions, as illustrated in Figure \(5\), it acts as a countercurrent multiplier. The vasa recta around it acts as the countercurrent exchanger.
Art Connection
Loop diuretics are drugs sometimes used to treat hypertension. These drugs inhibit the reabsorption of Na+ and Cl- ions by the ascending limb of the loop of Henle. A side effect is that they increase urination. Why do you think this is the case?
By the time the filtrate reaches the DCT, most of the urine and solutes have been reabsorbed. If the body requires additional water, all of it can be reabsorbed at this point. Further reabsorption is controlled by hormones, which will be discussed in a later section. Excretion of wastes occurs due to lack of reabsorption combined with tubular secretion. Undesirable products like metabolic wastes, urea, uric acid, and certain drugs, are excreted by tubular secretion. Most of the tubular secretion happens in the DCT, but some occurs in the early part of the collecting duct. Kidneys also maintain an acid-base balance by secreting excess H+ ions.
Although parts of the renal tubules are named proximal and distal, in a cross-section of the kidney, the tubules are placed close together and in contact with each other and the glomerulus. This allows for exchange of chemical messengers between the different cell types. For example, the DCT ascending limb of the loop of Henle has masses of cells called macula densa, which are in contact with cells of the afferent arterioles called juxtaglomerular cells. Together, the macula densa and juxtaglomerular cells form the juxtaglomerular complex (JGC). The JGC is an endocrine structure that secretes the enzyme renin and the hormone erythropoietin. When hormones trigger the macula densa cells in the DCT due to variations in blood volume, blood pressure, or electrolyte balance, these cells can immediately communicate the problem to the capillaries in the afferent and efferent arterioles, which can constrict or relax to change the glomerular filtration rate of the kidneys.
Career Connection: Nephrologist
A nephrologist studies and deals with diseases of the kidneys—both those that cause kidney failure (such as diabetes) and the conditions that are produced by kidney disease (such as hypertension). Blood pressure, blood volume, and changes in electrolyte balance come under the purview of a nephrologist.
Nephrologists usually work with other physicians who refer patients to them or consult with them about specific diagnoses and treatment plans. Patients are usually referred to a nephrologist for symptoms such as blood or protein in the urine, very high blood pressure, kidney stones, or renal failure.
Nephrology is a subspecialty of internal medicine. To become a nephrologist, medical school is followed by additional training to become certified in internal medicine. An additional two or more years is spent specifically studying kidney disorders and their accompanying effects on the body.
Summary
The kidneys are the main osmoregulatory organs in mammalian systems; they function to filter blood and maintain the osmolarity of body fluids at 300 mOsm. They are surrounded by three layers and are made up internally of three distinct regions—the cortex, medulla, and pelvis.
The blood vessels that transport blood into and out of the kidneys arise from and merge with the aorta and inferior vena cava, respectively. The renal arteries branch out from the aorta and enter the kidney where they further divide into segmental, interlobar, arcuate, and cortical radiate arteries.
The nephron is the functional unit of the kidney, which actively filters blood and generates urine. The nephron is made up of the renal corpuscle and renal tubule. Cortical nephrons are found in the renal cortex, while juxtamedullary nephrons are found in the renal cortex close to the renal medulla. The nephron filters and exchanges water and solutes with two sets of blood vessels and the tissue fluid in the kidneys.
There are three steps in the formation of urine: glomerular filtration, which occurs in the glomerulus; tubular reabsorption, which occurs in the renal tubules; and tubular secretion, which also occurs in the renal tubules.
Art Connections
Figure \(2\): Which of the following statements about the kidney is false?
1. The renal pelvis drains into the ureter.
2. The renal pyramids are in the medulla.
3. The cortex covers the capsule.
4. Nephrons are in the renal cortex.
Answer
C
Figure \(3\): Which of the following statements about the nephron is false?
1. The collecting duct empties into the distal convoluted tubule.
2. The Bowman’s capsule surrounds the glomerulus.
3. The loop of Henle is between the proximal and distal convoluted tubules.
4. The loop of Henle empties into the distal convoluted tubule.
Answer
A
Figure \(5\): Loop diuretics are drugs sometimes used to treat hypertension. These drugs inhibit the reabsorption of Na+ and Cl- ions by the ascending limb of the loop of Henle. A side effect is that they increase urination. Why do you think this is the case?
Answer
Loop diuretics decrease the excretion of salt into the renal medulla, thereby reducing its osmolality. As a result, less water is excreted into the medulla by the descending limb, and more water is excreted as urine.
Glossary
afferent arteriole
arteriole that branches from the cortical radiate artery and enters the glomerulus
arcuate artery
artery that branches from the interlobar artery and arches over the base of the renal pyramids
ascending limb
part of the loop of Henle that ascends from the renal medulla to the renal cortex
Bowman's capsule
structure that encloses the glomerulus
calyx
structure that connects the renal pelvis to the renal medulla
cortex (animal)
outer layer of an organ like the kidney or adrenal gland
cortical nephron
nephron that lies in the renal cortex
cortical radiate artery
artery that radiates from the arcuate arteries into the renal cortex
countercurrent exchanger
peritubular capillary network that allows exchange of solutes and water from the renal tubules
countercurrent multiplier
osmotic gradient in the renal medulla that is responsible for concentration of urine
descending limb
part of the loop of Henle that descends from the renal cortex into the renal medulla
distal convoluted tubule (DCT)
part of the renal tubule that is the most distant from the glomerulus
efferent arteriole
arteriole that exits from the glomerulus
glomerular filtration
filtration of blood in the glomerular capillary network into the glomerulus
glomerular filtration rate (GFR)
amount of filtrate formed by the glomerulus per minute
glomerulus (renal)
part of the renal corpuscle that contains the capillary network
hilum
region in the renal pelvis where blood vessels, nerves, and ureters bunch before entering or exiting the kidney
inferior vena cava
one of the main veins in the human body
interlobar artery
artery that branches from the segmental artery and travels in between the renal lobes
juxtaglomerular cell
cell in the afferent and efferent arterioles that responds to stimuli from the macula densa
juxtamedullary nephron
nephron that lies in the cortex but close to the renal medulla
kidney
organ that performs excretory and osmoregulatory functions
lobes of the kidney
renal pyramid along with the adjoining cortical region
loop of Henle
part of the renal tubule that loops into the renal medulla
macula densa
group of cells that senses changes in sodium ion concentration; present in parts of the renal tubule and collecting ducts
medulla
middle layer of an organ like the kidney or adrenal gland
nephron
functional unit of the kidney
perirenal fat capsule
fat layer that suspends the kidneys
peritubular capillary network
capillary network that surrounds the renal tubule after the efferent artery exits the glomerulus
proximal convoluted tubule (PCT)
part of the renal tubule that lies close to the glomerulus
renal artery
branch of the artery that enters the kidney
renal capsule
layer that encapsulates the kidneys
renal column
area of the kidney through which the interlobar arteries travel in the process of supplying blood to the renal lobes
renal corpuscle
glomerulus and the Bowman's capsule together
renal fascia
connective tissue that supports the kidneys
renal pelvis
region in the kidney where the calyces join the ureters
renal pyramid
conical structure in the renal medulla
renal tubule
tubule of the nephron that arises from the glomerulus
renal vein
branch of a vein that exits the kidney and joins the inferior vena cava
segmental artery
artery that branches from the renal artery
transport maximum
maximum amount of solute that can be transported out of the renal tubules during reabsorption
tubular reabsorption
reclamation of water and solutes that got filtered out in the glomerulus
tubular secretion
process of secretion of wastes that do not get reabsorbed
ureter
urine-bearing tube coming out of the kidney; carries urine to the bladder
urinary bladder
structure that the ureters empty the urine into; stores urine
urine
filtrate produced by kidneys that gets excreted out of the body
vasa recta
peritubular network that surrounds the loop of Henle of the juxtamedullary nephrons | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/49%3A_Osmotic_Regulation_and_the_Urinary_System/49.03%3A_Osmoregulatory_Organs.txt |
Skills to Develop
• Explain how the kidneys serve as the main osmoregulatory organs in mammalian systems
• Describe the structure of the kidneys and the functions of the parts of the kidney
• Describe how the nephron is the functional unit of the kidney and explain how it actively filters blood and generates urine
• Detail the three steps in the formation of urine: glomerular filtration, tubular reabsorption, and tubular secretion
Although the kidneys are the major osmoregulatory organ, the skin and lungs also play a role in the process. Water and electrolytes are lost through sweat glands in the skin, which helps moisturize and cool the skin surface, while the lungs expel a small amount of water in the form of mucous secretions and via evaporation of water vapor.
Kidneys: The Main Osmoregulatory Organ
The kidneys, illustrated in Figure \(1\), are a pair of bean-shaped structures that are located just below and posterior to the liver in the peritoneal cavity. The adrenal glands sit on top of each kidney and are also called the suprarenal glands. Kidneys filter blood and purify it. All the blood in the human body is filtered many times a day by the kidneys; these organs use up almost 25 percent of the oxygen absorbed through the lungs to perform this function. Oxygen allows the kidney cells to efficiently manufacture chemical energy in the form of ATP through aerobic respiration. The filtrate coming out of the kidneys is called urine.
Kidney Structure
Externally, the kidneys are surrounded by three layers, illustrated in Figure \(2\). The outermost layer is a tough connective tissue layer called the renal fascia. The second layer is called the perirenal fat capsule, which helps anchor the kidneys in place. The third and innermost layer is the renal capsule. Internally, the kidney has three regions—an outer cortex, a medulla in the middle, and the renal pelvis in the region called the hilum of the kidney. The hilum is the concave part of the bean-shape where blood vessels and nerves enter and exit the kidney; it is also the point of exit for the ureters. The renal cortex is granular due to the presence of nephrons—the functional unit of the kidney. The medulla consists of multiple pyramidal tissue masses, called the renal pyramids. In between the pyramids are spaces called renal columns through which the blood vessels pass. The tips of the pyramids, called renal papillae, point toward the renal pelvis. There are, on average, eight renal pyramids in each kidney. The renal pyramids along with the adjoining cortical region are called the lobes of the kidney. The renal pelvis leads to the ureter on the outside of the kidney. On the inside of the kidney, the renal pelvis branches out into two or three extensions called the major calyces, which further branch into the minor calyces. The ureters are urine-bearing tubes that exit the kidney and empty into the urinary bladder.
Art Connection
Which of the following statements about the kidney is false?
1. The renal pelvis drains into the ureter.
2. The renal pyramids are in the medulla.
3. The cortex covers the capsule.
4. Nephrons are in the renal cortex.
Because the kidney filters blood, its network of blood vessels is an important component of its structure and function. The arteries, veins, and nerves that supply the kidney enter and exit at the renal hilum. Renal blood supply starts with the branching of the aorta into the renal arteries (which are each named based on the region of the kidney they pass through) and ends with the exiting of the renal veins to join the inferior vena cava. The renal arteries split into several segmental arteries upon entering the kidneys. Each segmental artery splits further into several interlobar arteries and enters the renal columns, which supply the renal lobes. The interlobar arteries split at the junction of the renal cortex and medulla to form the arcuate arteries. The arcuate “bow shaped” arteries form arcs along the base of the medullary pyramids. Cortical radiate arteries, as the name suggests, radiate out from the arcuate arteries. The cortical radiate arteries branch into numerous afferent arterioles, and then enter the capillaries supplying the nephrons. Veins trace the path of the arteries and have similar names, except there are no segmental veins.
As mentioned previously, the functional unit of the kidney is the nephron, illustrated in Figure \(3\). Each kidney is made up of over one million nephrons that dot the renal cortex, giving it a granular appearance when sectioned sagittally. There are two types of nephrons—cortical nephrons (85 percent), which are deep in the renal cortex, and juxtamedullary nephrons (15 percent), which lie in the renal cortex close to the renal medulla. A nephron consists of three parts—a renal corpuscle, a renal tubule, and the associated capillary network, which originates from the cortical radiate arteries.
Art Connection
Which of the following statements about the nephron is false?
1. The collecting duct empties into the distal convoluted tubule.
2. The Bowman’s capsule surrounds the glomerulus.
3. The loop of Henle is between the proximal and distal convoluted tubules.
4. The loop of Henle empties into the distal convoluted tubule.
Renal Corpuscle
The renal corpuscle, located in the renal cortex, is made up of a network of capillaries known as the glomerulus and the capsule, a cup-shaped chamber that surrounds it, called the glomerular or Bowman's capsule.
Renal Tubule
The renal tubule is a long and convoluted structure that emerges from the glomerulus and can be divided into three parts based on function. The first part is called the proximal convoluted tubule (PCT) due to its proximity to the glomerulus; it stays in the renal cortex. The second part is called the loop of Henle, or nephritic loop, because it forms a loop (with descending and ascending limbs) that goes through the renal medulla. The third part of the renal tubule is called the distal convoluted tubule (DCT) and this part is also restricted to the renal cortex. The DCT, which is the last part of the nephron, connects and empties its contents into collecting ducts that line the medullary pyramids. The collecting ducts amass contents from multiple nephrons and fuse together as they enter the papillae of the renal medulla.
Capillary Network within the Nephron
The capillary network that originates from the renal arteries supplies the nephron with blood that needs to be filtered. The branch that enters the glomerulus is called the afferent arteriole. The branch that exits the glomerulus is called the efferent arteriole. Within the glomerulus, the network of capillaries is called the glomerular capillary bed. Once the efferent arteriole exits the glomerulus, it forms the peritubular capillary network, which surrounds and interacts with parts of the renal tubule. In cortical nephrons, the peritubular capillary network surrounds the PCT and DCT. In juxtamedullary nephrons, the peritubular capillary network forms a network around the loop of Henle and is called the vasa recta.
Kidney Function and Physiology
Kidneys filter blood in a three-step process. First, the nephrons filter blood that runs through the capillary network in the glomerulus. Almost all solutes, except for proteins, are filtered out into the glomerulus by a process called glomerular filtration. Second, the filtrate is collected in the renal tubules. Most of the solutes get reabsorbed in the PCT by a process called tubular reabsorption. In the loop of Henle, the filtrate continues to exchange solutes and water with the renal medulla and the peritubular capillary network. Water is also reabsorbed during this step. Then, additional solutes and wastes are secreted into the kidney tubules during tubular secretion, which is, in essence, the opposite process to tubular reabsorption. The collecting ducts collect filtrate coming from the nephrons and fuse in the medullary papillae. From here, the papillae deliver the filtrate, now called urine, into the minor calyces that eventually connect to the ureters through the renal pelvis. This entire process is illustrated in Figure \(4\).
Glomerular Filtration
Glomerular filtration filters out most of the solutes due to high blood pressure and specialized membranes in the afferent arteriole. The blood pressure in the glomerulus is maintained independent of factors that affect systemic blood pressure. The “leaky” connections between the endothelial cells of the glomerular capillary network allow solutes to pass through easily. All solutes in the glomerular capillaries, except for macromolecules like proteins, pass through by passive diffusion. There is no energy requirement at this stage of the filtration process. Glomerular filtration rate (GFR) is the volume of glomerular filtrate formed per minute by the kidneys. GFR is regulated by multiple mechanisms and is an important indicator of kidney function.
Link to Learning
To learn more about the vascular system of kidneys, click through this review and the steps of blood flow.
Tubular Reabsorption and Secretion
Tubular reabsorption occurs in the PCT part of the renal tubule. Almost all nutrients are reabsorbed, and this occurs either by passive or active transport. Reabsorption of water and some key electrolytes are regulated and can be influenced by hormones. Sodium (Na+) is the most abundant ion and most of it is reabsorbed by active transport and then transported to the peritubular capillaries. Because Na+ is actively transported out of the tubule, water follows it to even out the osmotic pressure. Water is also independently reabsorbed into the peritubular capillaries due to the presence of aquaporins, or water channels, in the PCT. This occurs due to the low blood pressure and high osmotic pressure in the peritubular capillaries. However, every solute has a transport maximum and the excess is not reabsorbed.
In the loop of Henle, the permeability of the membrane changes. The descending limb is permeable to water, not solutes; the opposite is true for the ascending limb. Additionally, the loop of Henle invades the renal medulla, which is naturally high in salt concentration and tends to absorb water from the renal tubule and concentrate the filtrate. The osmotic gradient increases as it moves deeper into the medulla. Because two sides of the loop of Henle perform opposing functions, as illustrated in Figure \(5\), it acts as a countercurrent multiplier. The vasa recta around it acts as the countercurrent exchanger.
Art Connection
Loop diuretics are drugs sometimes used to treat hypertension. These drugs inhibit the reabsorption of Na+ and Cl- ions by the ascending limb of the loop of Henle. A side effect is that they increase urination. Why do you think this is the case?
By the time the filtrate reaches the DCT, most of the urine and solutes have been reabsorbed. If the body requires additional water, all of it can be reabsorbed at this point. Further reabsorption is controlled by hormones, which will be discussed in a later section. Excretion of wastes occurs due to lack of reabsorption combined with tubular secretion. Undesirable products like metabolic wastes, urea, uric acid, and certain drugs, are excreted by tubular secretion. Most of the tubular secretion happens in the DCT, but some occurs in the early part of the collecting duct. Kidneys also maintain an acid-base balance by secreting excess H+ ions.
Although parts of the renal tubules are named proximal and distal, in a cross-section of the kidney, the tubules are placed close together and in contact with each other and the glomerulus. This allows for exchange of chemical messengers between the different cell types. For example, the DCT ascending limb of the loop of Henle has masses of cells called macula densa, which are in contact with cells of the afferent arterioles called juxtaglomerular cells. Together, the macula densa and juxtaglomerular cells form the juxtaglomerular complex (JGC). The JGC is an endocrine structure that secretes the enzyme renin and the hormone erythropoietin. When hormones trigger the macula densa cells in the DCT due to variations in blood volume, blood pressure, or electrolyte balance, these cells can immediately communicate the problem to the capillaries in the afferent and efferent arterioles, which can constrict or relax to change the glomerular filtration rate of the kidneys.
Career Connection: Nephrologist
A nephrologist studies and deals with diseases of the kidneys—both those that cause kidney failure (such as diabetes) and the conditions that are produced by kidney disease (such as hypertension). Blood pressure, blood volume, and changes in electrolyte balance come under the purview of a nephrologist.
Nephrologists usually work with other physicians who refer patients to them or consult with them about specific diagnoses and treatment plans. Patients are usually referred to a nephrologist for symptoms such as blood or protein in the urine, very high blood pressure, kidney stones, or renal failure.
Nephrology is a subspecialty of internal medicine. To become a nephrologist, medical school is followed by additional training to become certified in internal medicine. An additional two or more years is spent specifically studying kidney disorders and their accompanying effects on the body.
Summary
The kidneys are the main osmoregulatory organs in mammalian systems; they function to filter blood and maintain the osmolarity of body fluids at 300 mOsm. They are surrounded by three layers and are made up internally of three distinct regions—the cortex, medulla, and pelvis.
The blood vessels that transport blood into and out of the kidneys arise from and merge with the aorta and inferior vena cava, respectively. The renal arteries branch out from the aorta and enter the kidney where they further divide into segmental, interlobar, arcuate, and cortical radiate arteries.
The nephron is the functional unit of the kidney, which actively filters blood and generates urine. The nephron is made up of the renal corpuscle and renal tubule. Cortical nephrons are found in the renal cortex, while juxtamedullary nephrons are found in the renal cortex close to the renal medulla. The nephron filters and exchanges water and solutes with two sets of blood vessels and the tissue fluid in the kidneys.
There are three steps in the formation of urine: glomerular filtration, which occurs in the glomerulus; tubular reabsorption, which occurs in the renal tubules; and tubular secretion, which also occurs in the renal tubules.
Art Connections
Figure \(2\): Which of the following statements about the kidney is false?
1. The renal pelvis drains into the ureter.
2. The renal pyramids are in the medulla.
3. The cortex covers the capsule.
4. Nephrons are in the renal cortex.
Answer
C
Figure \(3\): Which of the following statements about the nephron is false?
1. The collecting duct empties into the distal convoluted tubule.
2. The Bowman’s capsule surrounds the glomerulus.
3. The loop of Henle is between the proximal and distal convoluted tubules.
4. The loop of Henle empties into the distal convoluted tubule.
Answer
A
Figure \(5\): Loop diuretics are drugs sometimes used to treat hypertension. These drugs inhibit the reabsorption of Na+ and Cl- ions by the ascending limb of the loop of Henle. A side effect is that they increase urination. Why do you think this is the case?
Answer
Loop diuretics decrease the excretion of salt into the renal medulla, thereby reducing its osmolality. As a result, less water is excreted into the medulla by the descending limb, and more water is excreted as urine.
Glossary
afferent arteriole
arteriole that branches from the cortical radiate artery and enters the glomerulus
arcuate artery
artery that branches from the interlobar artery and arches over the base of the renal pyramids
ascending limb
part of the loop of Henle that ascends from the renal medulla to the renal cortex
Bowman's capsule
structure that encloses the glomerulus
calyx
structure that connects the renal pelvis to the renal medulla
cortex (animal)
outer layer of an organ like the kidney or adrenal gland
cortical nephron
nephron that lies in the renal cortex
cortical radiate artery
artery that radiates from the arcuate arteries into the renal cortex
countercurrent exchanger
peritubular capillary network that allows exchange of solutes and water from the renal tubules
countercurrent multiplier
osmotic gradient in the renal medulla that is responsible for concentration of urine
descending limb
part of the loop of Henle that descends from the renal cortex into the renal medulla
distal convoluted tubule (DCT)
part of the renal tubule that is the most distant from the glomerulus
efferent arteriole
arteriole that exits from the glomerulus
glomerular filtration
filtration of blood in the glomerular capillary network into the glomerulus
glomerular filtration rate (GFR)
amount of filtrate formed by the glomerulus per minute
glomerulus (renal)
part of the renal corpuscle that contains the capillary network
hilum
region in the renal pelvis where blood vessels, nerves, and ureters bunch before entering or exiting the kidney
inferior vena cava
one of the main veins in the human body
interlobar artery
artery that branches from the segmental artery and travels in between the renal lobes
juxtaglomerular cell
cell in the afferent and efferent arterioles that responds to stimuli from the macula densa
juxtamedullary nephron
nephron that lies in the cortex but close to the renal medulla
kidney
organ that performs excretory and osmoregulatory functions
lobes of the kidney
renal pyramid along with the adjoining cortical region
loop of Henle
part of the renal tubule that loops into the renal medulla
macula densa
group of cells that senses changes in sodium ion concentration; present in parts of the renal tubule and collecting ducts
medulla
middle layer of an organ like the kidney or adrenal gland
nephron
functional unit of the kidney
perirenal fat capsule
fat layer that suspends the kidneys
peritubular capillary network
capillary network that surrounds the renal tubule after the efferent artery exits the glomerulus
proximal convoluted tubule (PCT)
part of the renal tubule that lies close to the glomerulus
renal artery
branch of the artery that enters the kidney
renal capsule
layer that encapsulates the kidneys
renal column
area of the kidney through which the interlobar arteries travel in the process of supplying blood to the renal lobes
renal corpuscle
glomerulus and the Bowman's capsule together
renal fascia
connective tissue that supports the kidneys
renal pelvis
region in the kidney where the calyces join the ureters
renal pyramid
conical structure in the renal medulla
renal tubule
tubule of the nephron that arises from the glomerulus
renal vein
branch of a vein that exits the kidney and joins the inferior vena cava
segmental artery
artery that branches from the renal artery
transport maximum
maximum amount of solute that can be transported out of the renal tubules during reabsorption
tubular reabsorption
reclamation of water and solutes that got filtered out in the glomerulus
tubular secretion
process of secretion of wastes that do not get reabsorbed
ureter
urine-bearing tube coming out of the kidney; carries urine to the bladder
urinary bladder
structure that the ureters empty the urine into; stores urine
urine
filtrate produced by kidneys that gets excreted out of the body
vasa recta
peritubular network that surrounds the loop of Henle of the juxtamedullary nephrons | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/49%3A_Osmotic_Regulation_and_the_Urinary_System/49.05%3A_The_Mammalian_Kidney.txt |
Skills to Develop
• Explain how hormonal cues help the kidneys synchronize the osmotic needs of the body
• Describe how hormones like epinephrine, norepinephrine, renin-angiotensin, aldosterone, anti-diuretic hormone, and atrial natriuretic peptide help regulate waste elimination, maintain correct osmolarity, and perform other osmoregulatory functions
While the kidneys operate to maintain osmotic balance and blood pressure in the body, they also act in concert with hormones. Hormones are small molecules that act as messengers within the body. Hormones are typically secreted from one cell and travel in the bloodstream to affect a target cell in another portion of the body. Different regions of the nephron bear specialized cells that have receptors to respond to chemical messengers and hormones. Table \(1\) summarizes the hormones that control the osmoregulatory functions.
Table \(1\): Hormones That Affect Osmoregulation
Hormone Where produced Function
Epinephrine and Norepinephrine Adrenal medulla Can decrease kidney function temporarily by vasoconstriction
Renin Kidney nephrons Increases blood pressure by acting on angiotensinogen
Angiotensin Liver Angiotensin II affects multiple processes and increases blood pressure
Aldosterone Adrenal cortex Prevents loss of sodium and water
Anti-diuretic hormone (vasopressin) Hypothalamus (stored in the posterior pituitary) Prevents water loss
Atrial natriuretic peptide Heart atrium Decreases blood pressure by acting as a vasodilator and increasing glomerular filtration rate; decreases sodium reabsorption in kidneys
Epinephrine and Norepinephrine
Epinephrine and norepinephrine are released by the adrenal medulla and nervous system respectively. They are the flight/fight hormones that are released when the body is under extreme stress. During stress, much of the body’s energy is used to combat imminent danger. Kidney function is halted temporarily by epinephrine and norepinephrine. These hormones function by acting directly on the smooth muscles of blood vessels to constrict them. Once the afferent arterioles are constricted, blood flow into the nephrons stops. These hormones go one step further and trigger the renin-angiotensin-aldosterone system.
Renin-Angiotensin-Aldosterone
The renin-angiotensin-aldosterone system, illustrated in Figure \(1\) proceeds through several steps to produce angiotensin II, which acts to stabilize blood pressure and volume. Renin (secreted by a part of the juxtaglomerular complex) is produced by the granular cells of the afferent and efferent arterioles. Thus, the kidneys control blood pressure and volume directly. Renin acts on angiotensinogen, which is made in the liver and converts it to angiotensin I. Angiotensin converting enzyme (ACE) converts angiotensin I to angiotensin II. Angiotensin II raises blood pressure by constricting blood vessels. It also triggers the release of the mineralocorticoid aldosterone from the adrenal cortex, which in turn stimulates the renal tubules to reabsorb more sodium. Angiotensin II also triggers the release of anti-diuretic hormone (ADH) from the hypothalamus, leading to water retention in the kidneys. It acts directly on the nephrons and decreases glomerular filtration rate. Medically, blood pressure can be controlled by drugs that inhibit ACE (called ACE inhibitors).
Mineralocorticoids
Mineralocorticoids are hormones synthesized by the adrenal cortex that affect osmotic balance. Aldosterone is a mineralocorticoid that regulates sodium levels in the blood. Almost all of the sodium in the blood is reclaimed by the renal tubules under the influence of aldosterone. Because sodium is always reabsorbed by active transport and water follows sodium to maintain osmotic balance, aldosterone manages not only sodium levels but also the water levels in body fluids. In contrast, the aldosterone also stimulates potassium secretion concurrently with sodium reabsorption. In contrast, absence of aldosterone means that no sodium gets reabsorbed in the renal tubules and all of it gets excreted in the urine. In addition, the daily dietary potassium load is not secreted and the retention of K+ can cause a dangerous increase in plasma K+ concentration. Patients who have Addison's disease have a failing adrenal cortex and cannot produce aldosterone. They lose sodium in their urine constantly, and if the supply is not replenished, the consequences can be fatal.
Antidiurectic Hormone
As previously discussed, antidiuretic hormone or ADH (also called vasopressin), as the name suggests, helps the body conserve water when body fluid volume, especially that of blood, is low. It is formed by the hypothalamus and is stored and released from the posterior pituitary. It acts by inserting aquaporins in the collecting ducts and promotes reabsorption of water. ADH also acts as a vasoconstrictor and increases blood pressure during hemorrhaging.
Atrial Natriuretic Peptide Hormone
The atrial natriuretic peptide (ANP) lowers blood pressure by acting as a vasodilator. It is released by cells in the atrium of the heart in response to high blood pressure and in patients with sleep apnea. ANP affects salt release, and because water passively follows salt to maintain osmotic balance, it also has a diuretic effect. ANP also prevents sodium reabsorption by the renal tubules, decreasing water reabsorption (thus acting as a diuretic) and lowering blood pressure. Its actions suppress the actions of aldosterone, ADH, and renin.
Summary
Hormonal cues help the kidneys synchronize the osmotic needs of the body. Hormones like epinephrine, norepinephrine, renin-angiotensin, aldosterone, anti-diuretic hormone, and atrial natriuretic peptide help regulate the needs of the body as well as the communication between the different organ systems.
Glossary
angiotensin converting enzyme (ACE)
enzyme that converts angiotensin I to angiotensin II
angiotensin I
product in the renin-angiotensin-aldosterone pathway
angiotensin II
molecule that affects different organs to increase blood pressure
anti-diuretic hormone (ADH)
hormone that prevents the loss of water
renin-angiotensin-aldosterone
biochemical pathway that activates angiotensin II, which increases blood pressure
vasodilator
compound that increases the diameter of blood vessels
vasopressin
another name for anti-diuretic hormone | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/49%3A_Osmotic_Regulation_and_the_Urinary_System/49.06%3A_Hormonal_Control_of_Osmoregulatory_Functions.txt |
• 50.1: Innate Immunity
The immune system comprises both innate and adaptive immune responses. Innate immunity occurs naturally because of genetic factors or physiology; it is not induced by infection or vaccination but works to reduce the workload for the adaptive immune response. Both the innate and adaptive levels of the immune response involve secreted proteins, receptor-mediated signaling, and intricate cell-to-cell communication.
• 50.2: Adaptive Immunity
The adaptive, or acquired, immune response takes days or even weeks to become established—much longer than the innate response; however, adaptive immunity is more specific to pathogens and has memory. Adaptive immunity is an immunity that occurs after exposure to an antigen either from a pathogen or a vaccination. This part of the immune system is activated when the innate immune response is insufficient to control an infection.
• 50.3: Cell-Mediated Immunity
The adaptive, or acquired, immune response takes days or even weeks to become established—much longer than the innate response; however, adaptive immunity is more specific to pathogens and has memory. Adaptive immunity is an immunity that occurs after exposure to an antigen either from a pathogen or a vaccination. This part of the immune system is activated when the innate immune response is insufficient to control an infection.
• 50.4: Humoral Immunity and Antibody Production
An antibody, also known as an immunoglobulin (Ig), is a protein that is produced by plasma cells after stimulation by an antigen. Antibodies are the functional basis of humoral immunity. Antibodies occur in the blood, in gastric and mucus secretions, and in breast milk. Antibodies in these bodily fluids can bind pathogens and mark them for destruction by phagocytes before they can infect cells.
• 50.5: Autoimmunity and Hypersensitivity
A functioning immune system is essential for survival, but even the sophisticated cellular and molecular defenses of the mammalian immune response can be defeated by pathogens at virtually every step. In the competition between immune protection and pathogen evasion, pathogens have the advantage of more rapid evolution because of their shorter generation time and other characteristics.
• 50.6: Antibodies in Medical Treatment and Diagnosis
• 50.7: Pathogens that Evade the Immune System
Viruses cause a variety of diseases in animals, including humans, ranging from the common cold to potentially fatal illnesses like meningitis . These diseases can be treated by antiviral drugs or by vaccines, but some viruses, such as HIV, are capable of both avoiding the immune response and mutating to become resistant to antiviral drugs.
50: The Immune System
Skills to Develop
• Describe physical and chemical immune barriers
• Explain immediate and induced innate immune responses
• Discuss natural killer cells
• Describe major histocompatibility class I molecules
• Summarize how the proteins in a complement system function to destroy extracellular pathogens
The immune system comprises both innate and adaptive immune responses. Innate immunity occurs naturally because of genetic factors or physiology; it is not induced by infection or vaccination but works to reduce the workload for the adaptive immune response. Both the innate and adaptive levels of the immune response involve secreted proteins, receptor-mediated signaling, and intricate cell-to-cell communication. The innate immune system developed early in animal evolution, roughly a billion years ago, as an essential response to infection. Innate immunity has a limited number of specific targets: any pathogenic threat triggers a consistent sequence of events that can identify the type of pathogen and either clear the infection independently or mobilize a highly specialized adaptive immune response. For example, tears and mucus secretions contain microbicidal factors.
Physical and Chemical Barriers
Before any immune factors are triggered, the skin functions as a continuous, impassable barrier to potentially infectious pathogens. Pathogens are killed or inactivated on the skin by desiccation (drying out) and by the skin’s acidity. In addition, beneficial microorganisms that coexist on the skin compete with invading pathogens, preventing infection. Regions of the body that are not protected by skin (such as the eyes and mucus membranes) have alternative methods of defense, such as tears and mucus secretions that trap and rinse away pathogens, and cilia in the nasal passages and respiratory tract that push the mucus with the pathogens out of the body. Throughout the body are other defenses, such as the low pH of the stomach (which inhibits the growth of pathogens), blood proteins that bind and disrupt bacterial cell membranes, and the process of urination (which flushes pathogens from the urinary tract).
Despite these barriers, pathogens may enter the body through skin abrasions or punctures, or by collecting on mucosal surfaces in large numbers that overcome the mucus or cilia. Some pathogens have evolved specific mechanisms that allow them to overcome physical and chemical barriers. When pathogens do enter the body, the innate immune system responds with inflammation, pathogen engulfment, and secretion of immune factors and proteins.
Pathogen Recognition
An infection may be intracellular or extracellular, depending on the pathogen. All viruses infect cells and replicate within those cells (intracellularly), whereas bacteria and other parasites may replicate intracellularly or extracellularly, depending on the species. The innate immune system must respond accordingly: by identifying the extracellular pathogen and/or by identifying host cells that have already been infected. When a pathogen enters the body, cells in the blood and lymph detect the specific pathogen-associated molecular patterns (PAMPs) on the pathogen’s surface. PAMPs are carbohydrate, polypeptide, and nucleic acid “signatures” that are expressed by viruses, bacteria, and parasites but which differ from molecules on host cells. The immune system has specific cells, described in Figure \(1\) and shown in Figure \(2\), with receptors that recognize these PAMPs. A macrophage is a large phagocytic cell that engulfs foreign particles and pathogens. Macrophages recognize PAMPs via complementary pattern recognition receptors (PRRs). PRRs are molecules on macrophages and dendritic cells which are in contact with the external environment. A monocyte is a type of white blood cell that circulates in the blood and lymph and differentiates into macrophages after it moves into infected tissue. Dendritic cells bind molecular signatures of pathogens and promote pathogen engulfment and destruction. Toll-like receptors (TLRs) are a type of PRR that recognizes molecules that are shared by pathogens but distinguishable from host molecules). TLRs are present in invertebrates as well as vertebrates, and appear to be one of the most ancient components of the immune system. TLRs have also been identified in the mammalian nervous system.
Cytokine Release Affect
The binding of PRRs with PAMPs triggers the release of cytokines, which signal that a pathogen is present and needs to be destroyed along with any infected cells. A cytokine is a chemical messenger that regulates cell differentiation (form and function), proliferation (production), and gene expression to affect immune responses. At least 40 types of cytokines exist in humans that differ in terms of the cell type that produces them, the cell type that responds to them, and the changes they produce. One type cytokine, interferon, is illustrated in Figure \(3\).
One subclass of cytokines is the interleukin (IL), so named because they mediate interactions between leukocytes (white blood cells). Interleukins are involved in bridging the innate and adaptive immune responses. In addition to being released from cells after PAMP recognition, cytokines are released by the infected cells which bind to nearby uninfected cells and induce those cells to release cytokines, which results in a cytokine burst.
A second class of early-acting cytokines is interferons, which are released by infected cells as a warning to nearby uninfected cells. One of the functions of an interferon is to inhibit viral replication. They also have other important functions, such as tumor surveillance. Interferons work by signaling neighboring uninfected cells to destroy RNA and reduce protein synthesis, signaling neighboring infected cells to undergo apoptosis (programmed cell death), and activating immune cells.
In response to interferons, uninfected cells alter their gene expression, which increases the cells’ resistance to infection. One effect of interferon-induced gene expression is a sharply reduced cellular protein synthesis. Virally infected cells produce more viruses by synthesizing large quantities of viral proteins. Thus, by reducing protein synthesis, a cell becomes resistant to viral infection.
Phagocytosis and Inflammation
The first cytokines to be produced are pro-inflammatory; that is, they encourage inflammation, the localized redness, swelling, heat, and pain that result from the movement of leukocytes and fluid through increasingly permeable capillaries to a site of infection. The population of leukocytes that arrives at an infection site depends on the nature of the infecting pathogen. Both macrophages and dendritic cells engulf pathogens and cellular debris through phagocytosis. A neutrophil is also a phagocytic leukocyte that engulfs and digests pathogens. Neutrophils, shown in Figure \(1\), are the most abundant leukocytes of the immune system. Neutrophils have a nucleus with two to five lobes, and they contain organelles, called lysosomes, that digest engulfed pathogens. An eosinophil is a leukocyte that works with other eosinophils to surround a parasite; it is involved in the allergic response and in protection against helminthes (parasitic worms).
Neutrophils and eosinophils are particularly important leukocytes that engulf large pathogens, such as bacteria and fungi. A mast cell is a leukocyte that produces inflammatory molecules, such as histamine, in response to large pathogens. A basophil is a leukocyte that, like a neutrophil, releases chemicals to stimulate the inflammatory response as illustrated in Figure \(4\). Basophils are also involved in allergy and hypersensitivity responses and induce specific types of inflammatory responses. Eosinophils and basophils produce additional inflammatory mediators to recruit more leukocytes. A hypersensitive immune response to harmless antigens, such as in pollen, often involves the release of histamine by basophils and mast cells.
Cytokines also send feedback to cells of the nervous system to bring about the overall symptoms of feeling sick, which include lethargy, muscle pain, and nausea. These effects may have evolved because the symptoms encourage the individual to rest and prevent them from spreading the infection to others. Cytokines also increase the core body temperature, causing a fever, which causes the liver to withhold iron from the blood. Without iron, certain pathogens, such as some bacteria, are unable to replicate; this is called nutritional immunity.
Natural Killer Cells
Lymphocytes are leukocytes that are histologically identifiable by their large, darkly staining nuclei; they are small cells with very little cytoplasm, as shown in Figure \(5\). Infected cells are identified and destroyed by natural killer (NK) cells, lymphocytes that can kill cells infected with viruses or tumor cells (abnormal cells that uncontrollably divide and invade other tissue). T cells and B cells of the adaptive immune system also are classified as lymphocytes. T cells are lymphocytes that mature in the thymus gland, and B cells are lymphocytes that mature in the bone marrow. NK cells identify intracellular infections, especially from viruses, by the altered expression of major histocompatibility class (MHC) I molecules on the surface of infected cells. MHC I molecules are proteins on the surfaces of all nucleated cells, thus they are scarce on red blood cells and platelets which are non-nucleated. The function of MHC I molecules is to display fragments of proteins from the infectious agents within the cell to T-cells; healthy cells will be ignored, while “non-self” or foreign proteins will be attacked by the immune system. MHC II molecules are found mainly on cells containing antigens (“non-self proteins”) and on lymphocytes. MHC II molecules interact with helper T-cells to trigger the appropriate immune response, which may include the inflammatory response.
An infected cell (or a tumor cell) is usually incapable of synthesizing and displaying MHC I molecules appropriately. The metabolic resources of cells infected by some viruses produce proteins that interfere with MHC I processing and/or trafficking to the cell surface. The reduced MHC I on host cells varies from virus to virus and results from active inhibitors being produced by the viruses. This process can deplete host MHC I molecules on the cell surface, which NK cells detect as “unhealthy” or “abnormal” while searching for cellular MHC I molecules. Similarly, the dramatically altered gene expression of tumor cells leads to expression of extremely deformed or absent MHC I molecules that also signal “unhealthy” or “abnormal.”
NK cells are always active; an interaction with normal, intact MHC I molecules on a healthy cell disables the killing sequence, and the NK cell moves on. After the NK cell detects an infected or tumor cell, its cytoplasm secretes granules comprised of perforin, a destructive protein that creates a pore in the target cell. Granzymes are released along with the perforin in the immunological synapse. A granzyme is a protease that digests cellular proteins and induces the target cell to undergo programmed cell death, or apoptosis. Phagocytic cells then digest the cell debris left behind. NK cells are constantly patrolling the body and are an effective mechanism for controlling potential infections and preventing cancer progression.
Complement
An array of approximately 20 types of soluble proteins, called a complement system, functions to destroy extracellular pathogens. Cells of the liver and macrophages synthesize complement proteins continuously; these proteins are abundant in the blood serum and are capable of responding immediately to infecting microorganisms. The complement system is so named because it is complementary to the antibody response of the adaptive immune system. Complement proteins bind to the surfaces of microorganisms and are particularly attracted to pathogens that are already bound by antibodies. Binding of complement proteins occurs in a specific and highly regulated sequence, with each successive protein being activated by cleavage and/or structural changes induced upon binding of the preceding protein(s). After the first few complement proteins bind, a cascade of sequential binding events follows in which the pathogen rapidly becomes coated in complement proteins.
Complement proteins perform several functions. The proteins serve as a marker to indicate the presence of a pathogen to phagocytic cells, such as macrophages and B cells, and enhance engulfment; this process is called opsonization. Certain complement proteins can combine to form attack complexes that open pores in microbial cell membranes. These structures destroy pathogens by causing their contents to leak, as illustrated in Figure \(6\).
Summary
The innate immune system serves as a first responder to pathogenic threats that bypass natural physical and chemical barriers of the body. Using a combination of cellular and molecular attacks, the innate immune system identifies the nature of a pathogen and responds with inflammation, phagocytosis, cytokine release, destruction by NK cells, and/or a complement system. When innate mechanisms are insufficient to clear an infection, the adaptive immune response is informed and mobilized.
Glossary
basophil
leukocyte that releases chemicals usually involved in the inflammatory response
B cell
lymphocyte that matures in the bone marrow and differentiates into antibody-secreting plasma cells
complement system
array of approximately 20 soluble proteins of the innate immune system that enhance phagocytosis, bore holes in pathogens, and recruit lymphocytes; enhances the adaptive response when antibodies are produced
cytokine
chemical messenger that regulates cell differentiation, proliferation, gene expression, and cell trafficking to effect immune responses
eosinophil
leukocyte that responds to parasites and is involved in the allergic response
granzyme
protease that enters target cells through perforin and induces apoptosis in the target cells; used by NK cells and killer T cells
inflammation
localized redness, swelling, heat, and pain that results from the movement of leukocytes and fluid through opened capillaries to a site of infection
innate immunity
immunity that occurs naturally because of genetic factors or physiology, and is not induced by infection or vaccination
interferon
cytokine that inhibits viral replication and modulates the immune response
lymphocyte
leukocyte that is histologically identifiable by its large nuclei; it is a small cell with very little cytoplasm
macrophage
large phagocytic cell that engulfs foreign particles and pathogens
major histocompatibility class (MHC) I/II molecule
protein found on the surface of all nucleated cells (I) or specifically on antigen-presenting cells (II) that signals to immune cells whether the cell is healthy/normal or is infected/cancerous; it provides the appropriate template into which antigens can be loaded for recognition by lymphocytes
mast cell
leukocyte that produces inflammatory molecules, such as histamine, in response to large pathogens and allergens
monocyte
type of white blood cell that circulates in the blood and lymph and differentiates into macrophages after it moves into infected tissue
natural killer (NK) cell
lymphocyte that can kill cells infected with viruses or tumor cells
neutrophil
phagocytic leukocyte that engulfs and digests pathogens
opsonization
process that enhances phagocytosis using proteins to indicate the presence of a pathogen to phagocytic cells
pathogen-associated molecular pattern (PAMP)
carbohydrate, polypeptide, and nucleic acid “signature” that is expressed by viruses, bacteria, and parasites but differs from molecules on host cells
pattern recognition receptor (PRR)
molecule on macrophages and dendritic cells that binds molecular signatures of pathogens and promotes pathogen engulfment and destruction
perforin
destructive protein that creates a pore in the target cell; used by NK cells and killer T cells
T cell
lymphocyte that matures in the thymus gland; one of the main cells involved in the adaptive immune system | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/50%3A_The_Immune_System/50.01%3A_Innate_Immunity.txt |
Skills to Develop
• Explain adaptive immunity
• Compare and contrast adaptive and innate immunity
• Describe cell-mediated immune response and humoral immune response
• Describe immune tolerance
The adaptive, or acquired, immune response takes days or even weeks to become established—much longer than the innate response; however, adaptive immunity is more specific to pathogens and has memory. Adaptive immunity is an immunity that occurs after exposure to an antigen either from a pathogen or a vaccination. This part of the immune system is activated when the innate immune response is insufficient to control an infection. In fact, without information from the innate immune system, the adaptive response could not be mobilized. There are two types of adaptive responses: the cell-mediated immune response, which is carried out by T cells, and the humoral immune response, which is controlled by activated B cells and antibodies. Activated T cells and B cells that are specific to molecular structures on the pathogen proliferate and attack the invading pathogen. Their attack can kill pathogens directly or secrete antibodies that enhance the phagocytosis of pathogens and disrupt the infection. Adaptive immunity also involves a memory to provide the host with long-term protection from reinfection with the same type of pathogen; on re-exposure, this memory will facilitate an efficient and quick response.
Antigen-presenting Cells
Unlike NK cells of the innate immune system, B cells (B lymphocytes) are a type of white blood cell that gives rise to antibodies, whereas T cells (T lymphocytes) are a type of white blood cell that plays an important role in the immune response. T cells are a key component in the cell-mediated response—the specific immune response that utilizes T cells to neutralize cells that have been infected with viruses and certain bacteria. There are three types of T cells: cytotoxic, helper, and suppressor T cells. Cytotoxic T cells destroy virus-infected cells in the cell-mediated immune response, and helper T cells play a part in activating both the antibody and the cell-mediated immune responses. Suppressor T cells deactivate T cells and B cells when needed, and thus prevent the immune response from becoming too intense.
An antigen is a foreign or “non-self” macromolecule that reacts with cells of the immune system. Not all antigens will provoke a response. For instance, individuals produce innumerable “self” antigens and are constantly exposed to harmless foreign antigens, such as food proteins, pollen, or dust components. The suppression of immune responses to harmless macromolecules is highly regulated and typically prevents processes that could be damaging to the host, known as tolerance.
The innate immune system contains cells that detect potentially harmful antigens, and then inform the adaptive immune response about the presence of these antigens. An antigen-presenting cell (APC) is an immune cell that detects, engulfs, and informs the adaptive immune response about an infection. When a pathogen is detected, these APCs will phagocytose the pathogen and digest it to form many different fragments of the antigen. Antigen fragments will then be transported to the surface of the APC, where they will serve as an indicator to other immune cells. Dendritic cells are immune cells that process antigen material; they are present in the skin (Langerhans cells) and the lining of the nose, lungs, stomach, and intestines. Sometimes a dendritic cell presents on the surface of other cells to induce an immune response, thus functioning as an antigen-presenting cell. Macrophages also function as APCs. Before activation and differentiation, B cells can also function as APCs.
After phagocytosis by APCs, the phagocytic vesicle fuses with an intracellular lysosome forming phagolysosome. Within the phagolysosome, the components are broken down into fragments; the fragments are then loaded onto MHC class I or MHC class II molecules and are transported to the cell surface for antigen presentation, as illustrated in Figure \(1\). Note that T lymphocytes cannot properly respond to the antigen unless it is processed and embedded in an MHC II molecule. APCs express MHC on their surfaces, and when combined with a foreign antigen, these complexes signal a “non-self” invader. Once the fragment of antigen is embedded in the MHC II molecule, the immune cell can respond. Helper T- cells are one of the main lymphocytes that respond to antigen-presenting cells. Recall that all other nucleated cells of the body expressed MHC I molecules, which signal “healthy” or “normal.”
T and B Lymphocytes
Lymphocytes in human circulating blood are approximately 80 to 90 percent T cells, shown in Figure \(2\), and 10 to 20 percent B cells. Recall that the T cells are involved in the cell-mediated immune response, whereas B cells are part of the humoral immune response.
T cells encompass a heterogeneous population of cells with extremely diverse functions. Some T cells respond to APCs of the innate immune system, and indirectly induce immune responses by releasing cytokines. Other T cells stimulate B cells to prepare their own response. Another population of T cells detects APC signals and directly kills the infected cells. Other T cells are involved in suppressing inappropriate immune reactions to harmless or “self” antigens.
T and B cells exhibit a common theme of recognition/binding of specific antigens via a complementary receptor, followed by activation and self-amplification/maturation to specifically bind to the particular antigen of the infecting pathogen. T and B lymphocytes are also similar in that each cell only expresses one type of antigen receptor. Any individual may possess a population of T and B cells that together express a near limitless variety of antigen receptors that are capable of recognizing virtually any infecting pathogen. T and B cells are activated when they recognize small components of antigens, called epitopes, presented by APCs, illustrated in Figure \(3\). Note that recognition occurs at a specific epitope rather than on the entire antigen; for this reason, epitopes are known as “antigenic determinants.” In the absence of information from APCs, T and B cells remain inactive, or naïve, and are unable to prepare an immune response. The requirement for information from the APCs of innate immunity to trigger B cell or T cell activation illustrates the essential nature of the innate immune response to the functioning of the entire immune system.
Naïve T cells can express one of two different molecules, CD4 or CD8, on their surface, as shown in Figure \(4\), and are accordingly classified as CD4+ or CD8+ cells. These molecules are important because they regulate how a T cell will interact with and respond to an APC. Naïve CD4+ cells bind APCs via their antigen-embedded MHC II molecules and are stimulated to become helper T (TH) lymphocytes, cells that go on to stimulate B cells (or cytotoxic T cells) directly or secrete cytokines to inform more and various target cells about the pathogenic threat. In contrast, CD8+ cells engage antigen-embedded MHC I molecules on APCs and are stimulated to become cytotoxic T lymphocytes (CTLs), which directly kill infected cells by apoptosis and emit cytokines to amplify the immune response. The two populations of T cells have different mechanisms of immune protection, but both bind MHC molecules via their antigen receptors called T cell receptors (TCRs). The CD4 or CD8 surface molecules differentiate whether the TCR will engage an MHC II or an MHC I molecule. Because they assist in binding specificity, the CD4 and CD8 molecules are described as coreceptors.
Exercise \(1\)
Which of the following statements about T cells is false?
1. Helper T cells release cytokines while cytotoxic T cells kill the infected cell.
2. Helper T cells are CD4+, while cytotoxic T cells are CD8+.
3. MHC II is a receptor found on most body cells, while MHC I is a receptor found on immune cells only.
4. The T cell receptor is found on both CD4+ and CD8+ T cells.
Answer
C
Consider the innumerable possible antigens that an individual will be exposed to during a lifetime. The mammalian adaptive immune system is adept in responding appropriately to each antigen. Mammals have an enormous diversity of T cell populations, resulting from the diversity of TCRs. Each TCR consists of two polypeptide chains that span the T cell membrane, as illustrated in Figure \(5\); the chains are linked by a disulfide bridge. Each polypeptide chain is comprised of a constant domain and a variable domain: a domain, in this sense, is a specific region of a protein that may be regulatory or structural. The intracellular domain is involved in intracellular signaling. A single T cell will express thousands of identical copies of one specific TCR variant on its cell surface. The specificity of the adaptive immune system occurs because it synthesizes millions of different T cell populations, each expressing a TCR that differs in its variable domain. This TCR diversity is achieved by the mutation and recombination of genes that encode these receptors in stem cell precursors of T cells. The binding between an antigen-displaying MHC molecule and a complementary TCR “match” indicates that the adaptive immune system needs to activate and produce that specific T cell because its structure is appropriate to recognize and destroy the invading pathogen.
Helper T Lymphocytes
The TH lymphocytes function indirectly to identify potential pathogens for other cells of the immune system. These cells are important for extracellular infections, such as those caused by certain bacteria, helminths, and protozoa. TH lymphocytes recognize specific antigens displayed in the MHC II complexes of APCs. There are two major populations of TH cells: TH1 and TH2. TH1 cells secrete cytokines to enhance the activities of macrophages and other T cells. TH1 cells activate the action of cyotoxic T cells, as well as macrophages. TH2 cells stimulate naïve B cells to destroy foreign invaders via antibody secretion. Whether a TH1 or a TH2 immune response develops depends on the specific types of cytokines secreted by cells of the innate immune system, which in turn depends on the nature of the invading pathogen.
The TH1-mediated response involves macrophages and is associated with inflammation. Recall the frontline defenses of macrophages involved in the innate immune response. Some intracellular bacteria, such as Mycobacterium tuberculosis, have evolved to multiply in macrophages after they have been engulfed. These pathogens evade attempts by macrophages to destroy and digest the pathogen. When M. tuberculosis infection occurs, macrophages can stimulate naïve T cells to become TH1 cells. These stimulated T cells secrete specific cytokines that send feedback to the macrophage to stimulate its digestive capabilities and allow it to destroy the colonizing M. tuberculosis. In the same manner, TH1-activated macrophages also become better suited to ingest and kill tumor cells. In summary; TH1 responses are directed toward intracellular invaders while TH2 responses are aimed at those that are extracellular.
B Lymphocytes
When stimulated by the TH2 pathway, naïve B cells differentiate into antibody-secreting plasma cells. A plasma cell is an immune cell that secrets antibodies; these cells arise from B cells that were stimulated by antigens. Similar to T cells, naïve B cells initially are coated in thousands of B cell receptors (BCRs), which are membrane-bound forms of Ig (immunoglobulin, or an antibody). The B cell receptor has two heavy chains and two light chains connected by disulfide linkages. Each chain has a constant and a variable region; the latter is involved in antigen binding. Two other membrane proteins, Ig alpha and Ig beta, are involved in signaling. The receptors of any particular B cell, as shown in Figure \(6\) are all the same, but the hundreds of millions of different B cells in an individual have distinct recognition domains that contribute to extensive diversity in the types of molecular structures to which they can bind. In this state, B cells function as APCs. They bind and engulf foreign antigens via their BCRs and then display processed antigens in the context of MHC II molecules to TH2 cells. When a TH2 cell detects that a B cell is bound to a relevant antigen, it secretes specific cytokines that induce the B cell to proliferate rapidly, which makes thousands of identical (clonal) copies of it, and then it synthesizes and secretes antibodies with the same antigen recognition pattern as the BCRs. The activation of B cells corresponding to one specific BCR variant and the dramatic proliferation of that variant is known as clonal selection. This phenomenon drastically, but briefly, changes the proportions of BCR variants expressed by the immune system, and shifts the balance toward BCRs specific to the infecting pathogen.
T and B cells differ in one fundamental way: whereas T cells bind antigens that have been digested and embedded in MHC molecules by APCs, B cells function as APCs that bind intact antigens that have not been processed. Although T and B cells both react with molecules that are termed “antigens,” these lymphocytes actually respond to very different types of molecules. B cells must be able to bind intact antigens because they secrete antibodies that must recognize the pathogen directly, rather than digested remnants of the pathogen. Bacterial carbohydrate and lipid molecules can activate B cells independently from the T cells.
Cytotoxic T Lymphocytes
CTLs, a subclass of T cells, function to clear infections directly. The cell-mediated part of the adaptive immune system consists of CTLs that attack and destroy infected cells. CTLs are particularly important in protecting against viral infections; this is because viruses replicate within cells where they are shielded from extracellular contact with circulating antibodies. When APCs phagocytize pathogens and present MHC I-embedded antigens to naïve CD8+ T cells that express complementary TCRs, the CD8+ T cells become activated to proliferate according to clonal selection. These resulting CTLs then identify non-APCs displaying the same MHC I-embedded antigens (for example, viral proteins)—for example, the CTLs identify infected host cells.
Intracellularly, infected cells typically die after the infecting pathogen replicates to a sufficient concentration and lyses the cell, as many viruses do. CTLs attempt to identify and destroy infected cells before the pathogen can replicate and escape, thereby halting the progression of intracellular infections. CTLs also support NK lymphocytes to destroy early cancers. Cytokines secreted by the TH1 response that stimulates macrophages also stimulate CTLs and enhance their ability to identify and destroy infected cells and tumors.
CTLs sense MHC I-embedded antigens by directly interacting with infected cells via their TCRs. Binding of TCRs with antigens activates CTLs to release perforin and granzyme, degradative enzymes that will induce apoptosis of the infected cell. Recall that this is a similar destruction mechanism to that used by NK cells. In this process, the CTL does not become infected and is not harmed by the secretion of perforin and granzymes. In fact, the functions of NK cells and CTLs are complementary and maximize the removal of infected cells, as illustrated in Figure \(7\). If the NK cell cannot identify the “missing self” pattern of down-regulated MHC I molecules, then the CTL can identify it by the complex of MHC I with foreign antigens, which signals “altered self.” Similarly, if the CTL cannot detect antigen-embedded MHC I because the receptors are depleted from the cell surface, NK cells will destroy the cell instead. CTLs also emit cytokines, such as interferons, that alter surface protein expression in other infected cells, such that the infected cells can be easily identified and destroyed. Moreover, these interferons can also prevent virally infected cells from releasing virus particles.
Exercise \(2\)
Based on what you know about MHC receptors, why do you think an organ transplanted from an incompatible donor to a recipient will be rejected?
Answer
MHC receptors differ from person to person. Thus, MHC receptors on an incompatible donor are considered “non-self” and are rejected by the immune system.
Plasma cells and CTLs are collectively called effector cells: they represent differentiated versions of their naïve counterparts, and they are involved in bringing about the immune defense of killing pathogens and infected host cells.
Mucosal Surfaces and Immune Tolerance
The innate and adaptive immune responses discussed thus far comprise the systemic immune system (affecting the whole body), which is distinct from the mucosal immune system. Mucosal immunity is formed by mucosa-associated lymphoid tissue, which functions independently of the systemic immune system, and which has its own innate and adaptive components. Mucosa-associated lymphoid tissue (MALT), illustrated in Figure \(8\), is a collection of lymphatic tissue that combines with epithelial tissue lining the mucosa throughout the body. This tissue functions as the immune barrier and response in areas of the body with direct contact to the external environment. The systemic and mucosal immune systems use many of the same cell types. Foreign particles that make their way to MALT are taken up by absorptive epithelial cells called M cells and delivered to APCs located directly below the mucosal tissue. M cells function in the transport described, and are located in the Peyer’s patch, a lymphoid nodule. APCs of the mucosal immune system are primarily dendritic cells, with B cells and macrophages having minor roles. Processed antigens displayed on APCs are detected by T cells in the MALT and at various mucosal induction sites, such as the tonsils, adenoids, appendix, or the mesenteric lymph nodes of the intestine. Activated T cells then migrate through the lymphatic system and into the circulatory system to mucosal sites of infection.
MALT is a crucial component of a functional immune system because mucosal surfaces, such as the nasal passages, are the first tissues onto which inhaled or ingested pathogens are deposited. The mucosal tissue includes the mouth, pharynx, and esophagus, and the gastrointestinal, respiratory, and urogenital tracts.
The immune system has to be regulated to prevent wasteful, unnecessary responses to harmless substances, and more importantly so that it does not attack “self.” The acquired ability to prevent an unnecessary or harmful immune response to a detected foreign substance known not to cause disease is described as immune tolerance. Immune tolerance is crucial for maintaining mucosal homeostasis given the tremendous number of foreign substances (such as food proteins) that APCs of the oral cavity, pharynx, and gastrointestinal mucosa encounter. Immune tolerance is brought about by specialized APCs in the liver, lymph nodes, small intestine, and lung that present harmless antigens to an exceptionally diverse population of regulatory T (Treg) cells, specialized lymphocytes that suppress local inflammation and inhibit the secretion of stimulatory immune factors. The combined result of Treg cells is to prevent immunologic activation and inflammation in undesired tissue compartments and to allow the immune system to focus on pathogens instead. In addition to promoting immune tolerance of harmless antigens, other subsets of Treg cells are involved in the prevention of the autoimmune response, which is an inappropriate immune response to host cells or self-antigens. Another Treg class suppresses immune responses to harmful pathogens after the infection has cleared to minimize host cell damage induced by inflammation and cell lysis.
Immunological Memory
The adaptive immune system possesses a memory component that allows for an efficient and dramatic response upon reinvasion of the same pathogen. Memory is handled by the adaptive immune system with little reliance on cues from the innate response. During the adaptive immune response to a pathogen that has not been encountered before, called a primary response, plasma cells secreting antibodies and differentiated T cells increase, then plateau over time. As B and T cells mature into effector cells, a subset of the naïve populations differentiates into B and T memory cells with the same antigen specificities, as illustrated in Figure \(9\).
A memory cell is an antigen-specific B or T lymphocyte that does not differentiate into effector cells during the primary immune response, but that can immediately become effector cells upon re-exposure to the same pathogen. During the primary immune response, memory cells do not respond to antigens and do not contribute to host defenses. As the infection is cleared and pathogenic stimuli subside, the effectors are no longer needed, and they undergo apoptosis. In contrast, the memory cells persist in the circulation.
Exercise \(3\)
The Rh antigen is found on Rh-positive red blood cells. An Rh-negative female can usually carry an Rh-positive fetus to term without difficulty. However, if she has a second Rh-positive fetus, her body may launch an immune attack that causes hemolytic disease of the newborn. Why do you think hemolytic disease is only a problem during the second or subsequent pregnancies?
Answer
If the blood of the mother and fetus mixes, memory cells that recognize the Rh antigen can form late in the first pregnancy. During subsequent pregnancies, these memory cells launch an immune attack on the fetal blood cells. Injection of anti-Rh antibody during the first pregnancy prevents the immune response from occurring.
If the pathogen is never encountered again during the individual’s lifetime, B and T memory cells will circulate for a few years or even several decades and will gradually die off, having never functioned as effector cells. However, if the host is re-exposed to the same pathogen type, circulating memory cells will immediately differentiate into plasma cells and CTLs without input from APCs or TH cells. One reason the adaptive immune response is delayed is because it takes time for naïve B and T cells with the appropriate antigen specificities to be identified and activated. Upon reinfection, this step is skipped, and the result is a more rapid production of immune defenses. Memory B cells that differentiate into plasma cells output tens to hundreds-fold greater antibody amounts than were secreted during the primary response, as the graph in Figure \(10\) illustrates. This rapid and dramatic antibody response may stop the infection before it can even become established, and the individual may not realize they had been exposed.
Vaccination is based on the knowledge that exposure to noninfectious antigens, derived from known pathogens, generates a mild primary immune response. The immune response to vaccination may not be perceived by the host as illness but still confers immune memory. When exposed to the corresponding pathogen to which an individual was vaccinated, the reaction is similar to a secondary exposure. Because each reinfection generates more memory cells and increased resistance to the pathogen, and because some memory cells die, certain vaccine courses involve one or more booster vaccinations to mimic repeat exposures: for instance, tetanus boosters are necessary every ten years because the memory cells only live that long.
Mucosal Immune Memory
A subset of T and B cells of the mucosal immune system differentiates into memory cells just as in the systemic immune system. Upon reinvasion of the same pathogen type, a pronounced immune response occurs at the mucosal site where the original pathogen deposited, but a collective defense is also organized within interconnected or adjacent mucosal tissue. For instance, the immune memory of an infection in the oral cavity would also elicit a response in the pharynx if the oral cavity was exposed to the same pathogen.
Career Connection: Vaccinologist
Vaccination (or immunization) involves the delivery, usually by injection as shown in Figure \(11\), of noninfectious antigen(s) derived from known pathogens. Other components, called adjuvants, are delivered in parallel to help stimulate the immune response. Immunological memory is the reason vaccines work. Ideally, the effect of vaccination is to elicit immunological memory, and thus resistance to specific pathogens without the individual having to experience an infection.
Vaccinologists are involved in the process of vaccine development from the initial idea to the availability of the completed vaccine. This process can take decades, can cost millions of dollars, and can involve many obstacles along the way. For instance, injected vaccines stimulate the systemic immune system, eliciting humoral and cell-mediated immunity, but have little effect on the mucosal response, which presents a challenge because many pathogens are deposited and replicate in mucosal compartments, and the injection does not provide the most efficient immune memory for these disease agents. For this reason, vaccinologists are actively involved in developing new vaccines that are applied via intranasal, aerosol, oral, or transcutaneous (absorbed through the skin) delivery methods. Importantly, mucosal-administered vaccines elicit both mucosal and systemic immunity and produce the same level of disease resistance as injected vaccines.
Currently, a version of intranasal influenza vaccine is available, and the polio and typhoid vaccines can be administered orally, as shown in Figure \(12\). Similarly, the measles and rubella vaccines are being adapted to aerosol delivery using inhalation devices. Eventually, transgenic plants may be engineered to produce vaccine antigens that can be eaten to confer disease resistance. Other vaccines may be adapted to rectal or vaginal application to elicit immune responses in rectal, genitourinary, or reproductive mucosa. Finally, vaccine antigens may be adapted to transdermal application in which the skin is lightly scraped and microneedles are used to pierce the outermost layer. In addition to mobilizing the mucosal immune response, this new generation of vaccines may end the anxiety associated with injections and, in turn, improve patient participation.
Primary Centers of the Immune System
Although the immune system is characterized by circulating cells throughout the body, the regulation, maturation, and intercommunication of immune factors occur at specific sites. The blood circulates immune cells, proteins, and other factors through the body. Approximately 0.1 percent of all cells in the blood are leukocytes, which encompass monocytes (the precursor of macrophages) and lymphocytes. The majority of cells in the blood are erythrocytes (red blood cells). Lymph is a watery fluid that bathes tissues and organs with protective white blood cells and does not contain erythrocytes. Cells of the immune system can travel between the distinct lymphatic and blood circulatory systems, which are separated by interstitial space, by a process called extravasation (passing through to surrounding tissue).
The cells of the immune system originate from hematopoietic stem cells in the bone marrow. Cytokines stimulate these stem cells to differentiate into immune cells. B cell maturation occurs in the bone marrow, whereas naïve T cells transit from the bone marrow to the thymus for maturation. In the thymus, immature T cells that express TCRs complementary to self-antigens are destroyed. This process helps prevent autoimmune responses.
On maturation, T and B lymphocytes circulate to various destinations. Lymph nodes scattered throughout the body, as illustrated in Figure \(13\), house large populations of T and B cells, dendritic cells, and macrophages. Lymph gathers antigens as it drains from tissues. These antigens then are filtered through lymph nodes before the lymph is returned to circulation. APCs in the lymph nodes capture and process antigens and inform nearby lymphocytes about potential pathogens.
The spleen houses B and T cells, macrophages, dendritic cells, and NK cells. The spleen, shown in Figure \(14\), is the site where APCs that have trapped foreign particles in the blood can communicate with lymphocytes. Antibodies are synthesized and secreted by activated plasma cells in the spleen, and the spleen filters foreign substances and antibody-complexed pathogens from the blood. Functionally, the spleen is to the blood as lymph nodes are to the lymph.
Summary
The adaptive immune response is a slower-acting, longer-lasting, and more specific response than the innate response. However, the adaptive response requires information from the innate immune system to function. APCs display antigens via MHC molecules to complementary naïve T cells. In response, the T cells differentiate and proliferate, becoming TH cells or CTLs. TH cells stimulate B cells that have engulfed and presented pathogen-derived antigens. B cells differentiate into plasma cells that secrete antibodies, whereas CTLs induce apoptosis in intracellularly infected or cancerous cells. Memory cells persist after a primary exposure to a pathogen. If re-exposure occurs, memory cells differentiate into effector cells without input from the innate immune system. The mucosal immune system is largely independent from the systemic immune system but functions in a parallel fashion to protect the extensive mucosal surfaces of the body.
Glossary
adaptive immunity
immunity that has memory and occurs after exposure to an antigen either from a pathogen or a vaccination
antigen
foreign or “non-self” protein that triggers the immune response
antigen-presenting cell (APC)
immune cell that detects, engulfs, and informs the adaptive immune response about an infection by presenting the processed antigen on the cell surface
autoimmune response
inappropriate immune response to host cells or self-antigens
cell-mediated immune response
adaptive immune response that is carried out by T cells
clonal selection
activation of B cells corresponding to one specific BCR variant and the dramatic proliferation of that variant
cytotoxic T lymphocyte (CTL)
adaptive immune cell that directly kills infected cells via perforin and granzymes, and releases cytokines to enhance the immune response
dendritic cell
immune cell that processes antigen material and presents it on the surface of other cells to induce an immune response
effector cell
lymphocyte that has differentiated, such as a B cell, plasma cell, or cytotoxic T lymphocyte
epitope
small component of an antigen that is specifically recognized by antibodies, B cells, and T cells; the antigenic determinant
helper T lymphocyte (TH)
cell of the adaptive immune system that binds APCs via MHC II molecules and stimulates B cells or secretes cytokines to initiate the immune response
humoral immune response
adaptive immune response that is controlled by activated B cells and antibodies
immune tolerance
acquired ability to prevent an unnecessary or harmful immune response to a detected foreign body known not to cause disease or to self-antigens
lymph
watery fluid that bathes tissues and organs with protective white blood cells and does not contain erythrocytes
mucosa-associated lymphoid tissue (MALT)
collection of lymphatic tissue that combines with epithelial tissue lining the mucosa throughout the body
memory cell
antigen-specific B or T lymphocyte that does not differentiate into effector cells during the primary immune response but that can immediately become an effector cell upon re-exposure to the same pathogen
plasma cell
immune cell that secrets antibodies; these cells arise from B cells that were stimulated by antigens
regulatory T (Treg) cell
specialized lymphocyte that suppresses local inflammation and inhibits the secretion of cytokines, antibodies, and other stimulatory immune factors; involved in immune tolerance | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/50%3A_The_Immune_System/50.02%3A_Adaptive_Immunity.txt |
Skills to Develop
• Explain adaptive immunity
• Compare and contrast adaptive and innate immunity
• Describe cell-mediated immune response and humoral immune response
• Describe immune tolerance
The adaptive, or acquired, immune response takes days or even weeks to become established—much longer than the innate response; however, adaptive immunity is more specific to pathogens and has memory. Adaptive immunity is an immunity that occurs after exposure to an antigen either from a pathogen or a vaccination. This part of the immune system is activated when the innate immune response is insufficient to control an infection. In fact, without information from the innate immune system, the adaptive response could not be mobilized. There are two types of adaptive responses: the cell-mediated immune response, which is carried out by T cells, and the humoral immune response, which is controlled by activated B cells and antibodies. Activated T cells and B cells that are specific to molecular structures on the pathogen proliferate and attack the invading pathogen. Their attack can kill pathogens directly or secrete antibodies that enhance the phagocytosis of pathogens and disrupt the infection. Adaptive immunity also involves a memory to provide the host with long-term protection from reinfection with the same type of pathogen; on re-exposure, this memory will facilitate an efficient and quick response.
Antigen-presenting Cells
Unlike NK cells of the innate immune system, B cells (B lymphocytes) are a type of white blood cell that gives rise to antibodies, whereas T cells (T lymphocytes) are a type of white blood cell that plays an important role in the immune response. T cells are a key component in the cell-mediated response—the specific immune response that utilizes T cells to neutralize cells that have been infected with viruses and certain bacteria. There are three types of T cells: cytotoxic, helper, and suppressor T cells. Cytotoxic T cells destroy virus-infected cells in the cell-mediated immune response, and helper T cells play a part in activating both the antibody and the cell-mediated immune responses. Suppressor T cells deactivate T cells and B cells when needed, and thus prevent the immune response from becoming too intense.
An antigen is a foreign or “non-self” macromolecule that reacts with cells of the immune system. Not all antigens will provoke a response. For instance, individuals produce innumerable “self” antigens and are constantly exposed to harmless foreign antigens, such as food proteins, pollen, or dust components. The suppression of immune responses to harmless macromolecules is highly regulated and typically prevents processes that could be damaging to the host, known as tolerance.
The innate immune system contains cells that detect potentially harmful antigens, and then inform the adaptive immune response about the presence of these antigens. An antigen-presenting cell (APC) is an immune cell that detects, engulfs, and informs the adaptive immune response about an infection. When a pathogen is detected, these APCs will phagocytose the pathogen and digest it to form many different fragments of the antigen. Antigen fragments will then be transported to the surface of the APC, where they will serve as an indicator to other immune cells. Dendritic cells are immune cells that process antigen material; they are present in the skin (Langerhans cells) and the lining of the nose, lungs, stomach, and intestines. Sometimes a dendritic cell presents on the surface of other cells to induce an immune response, thus functioning as an antigen-presenting cell. Macrophages also function as APCs. Before activation and differentiation, B cells can also function as APCs.
After phagocytosis by APCs, the phagocytic vesicle fuses with an intracellular lysosome forming phagolysosome. Within the phagolysosome, the components are broken down into fragments; the fragments are then loaded onto MHC class I or MHC class II molecules and are transported to the cell surface for antigen presentation, as illustrated in Figure \(1\). Note that T lymphocytes cannot properly respond to the antigen unless it is processed and embedded in an MHC II molecule. APCs express MHC on their surfaces, and when combined with a foreign antigen, these complexes signal a “non-self” invader. Once the fragment of antigen is embedded in the MHC II molecule, the immune cell can respond. Helper T- cells are one of the main lymphocytes that respond to antigen-presenting cells. Recall that all other nucleated cells of the body expressed MHC I molecules, which signal “healthy” or “normal.”
T and B Lymphocytes
Lymphocytes in human circulating blood are approximately 80 to 90 percent T cells, shown in Figure \(2\), and 10 to 20 percent B cells. Recall that the T cells are involved in the cell-mediated immune response, whereas B cells are part of the humoral immune response.
T cells encompass a heterogeneous population of cells with extremely diverse functions. Some T cells respond to APCs of the innate immune system, and indirectly induce immune responses by releasing cytokines. Other T cells stimulate B cells to prepare their own response. Another population of T cells detects APC signals and directly kills the infected cells. Other T cells are involved in suppressing inappropriate immune reactions to harmless or “self” antigens.
T and B cells exhibit a common theme of recognition/binding of specific antigens via a complementary receptor, followed by activation and self-amplification/maturation to specifically bind to the particular antigen of the infecting pathogen. T and B lymphocytes are also similar in that each cell only expresses one type of antigen receptor. Any individual may possess a population of T and B cells that together express a near limitless variety of antigen receptors that are capable of recognizing virtually any infecting pathogen. T and B cells are activated when they recognize small components of antigens, called epitopes, presented by APCs, illustrated in Figure \(3\). Note that recognition occurs at a specific epitope rather than on the entire antigen; for this reason, epitopes are known as “antigenic determinants.” In the absence of information from APCs, T and B cells remain inactive, or naïve, and are unable to prepare an immune response. The requirement for information from the APCs of innate immunity to trigger B cell or T cell activation illustrates the essential nature of the innate immune response to the functioning of the entire immune system.
Naïve T cells can express one of two different molecules, CD4 or CD8, on their surface, as shown in Figure \(4\), and are accordingly classified as CD4+ or CD8+ cells. These molecules are important because they regulate how a T cell will interact with and respond to an APC. Naïve CD4+ cells bind APCs via their antigen-embedded MHC II molecules and are stimulated to become helper T (TH) lymphocytes, cells that go on to stimulate B cells (or cytotoxic T cells) directly or secrete cytokines to inform more and various target cells about the pathogenic threat. In contrast, CD8+ cells engage antigen-embedded MHC I molecules on APCs and are stimulated to become cytotoxic T lymphocytes (CTLs), which directly kill infected cells by apoptosis and emit cytokines to amplify the immune response. The two populations of T cells have different mechanisms of immune protection, but both bind MHC molecules via their antigen receptors called T cell receptors (TCRs). The CD4 or CD8 surface molecules differentiate whether the TCR will engage an MHC II or an MHC I molecule. Because they assist in binding specificity, the CD4 and CD8 molecules are described as coreceptors.
Exercise \(1\)
Which of the following statements about T cells is false?
1. Helper T cells release cytokines while cytotoxic T cells kill the infected cell.
2. Helper T cells are CD4+, while cytotoxic T cells are CD8+.
3. MHC II is a receptor found on most body cells, while MHC I is a receptor found on immune cells only.
4. The T cell receptor is found on both CD4+ and CD8+ T cells.
Answer
C
Consider the innumerable possible antigens that an individual will be exposed to during a lifetime. The mammalian adaptive immune system is adept in responding appropriately to each antigen. Mammals have an enormous diversity of T cell populations, resulting from the diversity of TCRs. Each TCR consists of two polypeptide chains that span the T cell membrane, as illustrated in Figure \(5\); the chains are linked by a disulfide bridge. Each polypeptide chain is comprised of a constant domain and a variable domain: a domain, in this sense, is a specific region of a protein that may be regulatory or structural. The intracellular domain is involved in intracellular signaling. A single T cell will express thousands of identical copies of one specific TCR variant on its cell surface. The specificity of the adaptive immune system occurs because it synthesizes millions of different T cell populations, each expressing a TCR that differs in its variable domain. This TCR diversity is achieved by the mutation and recombination of genes that encode these receptors in stem cell precursors of T cells. The binding between an antigen-displaying MHC molecule and a complementary TCR “match” indicates that the adaptive immune system needs to activate and produce that specific T cell because its structure is appropriate to recognize and destroy the invading pathogen.
Helper T Lymphocytes
The TH lymphocytes function indirectly to identify potential pathogens for other cells of the immune system. These cells are important for extracellular infections, such as those caused by certain bacteria, helminths, and protozoa. TH lymphocytes recognize specific antigens displayed in the MHC II complexes of APCs. There are two major populations of TH cells: TH1 and TH2. TH1 cells secrete cytokines to enhance the activities of macrophages and other T cells. TH1 cells activate the action of cyotoxic T cells, as well as macrophages. TH2 cells stimulate naïve B cells to destroy foreign invaders via antibody secretion. Whether a TH1 or a TH2 immune response develops depends on the specific types of cytokines secreted by cells of the innate immune system, which in turn depends on the nature of the invading pathogen.
The TH1-mediated response involves macrophages and is associated with inflammation. Recall the frontline defenses of macrophages involved in the innate immune response. Some intracellular bacteria, such as Mycobacterium tuberculosis, have evolved to multiply in macrophages after they have been engulfed. These pathogens evade attempts by macrophages to destroy and digest the pathogen. When M. tuberculosis infection occurs, macrophages can stimulate naïve T cells to become TH1 cells. These stimulated T cells secrete specific cytokines that send feedback to the macrophage to stimulate its digestive capabilities and allow it to destroy the colonizing M. tuberculosis. In the same manner, TH1-activated macrophages also become better suited to ingest and kill tumor cells. In summary; TH1 responses are directed toward intracellular invaders while TH2 responses are aimed at those that are extracellular.
B Lymphocytes
When stimulated by the TH2 pathway, naïve B cells differentiate into antibody-secreting plasma cells. A plasma cell is an immune cell that secrets antibodies; these cells arise from B cells that were stimulated by antigens. Similar to T cells, naïve B cells initially are coated in thousands of B cell receptors (BCRs), which are membrane-bound forms of Ig (immunoglobulin, or an antibody). The B cell receptor has two heavy chains and two light chains connected by disulfide linkages. Each chain has a constant and a variable region; the latter is involved in antigen binding. Two other membrane proteins, Ig alpha and Ig beta, are involved in signaling. The receptors of any particular B cell, as shown in Figure \(6\) are all the same, but the hundreds of millions of different B cells in an individual have distinct recognition domains that contribute to extensive diversity in the types of molecular structures to which they can bind. In this state, B cells function as APCs. They bind and engulf foreign antigens via their BCRs and then display processed antigens in the context of MHC II molecules to TH2 cells. When a TH2 cell detects that a B cell is bound to a relevant antigen, it secretes specific cytokines that induce the B cell to proliferate rapidly, which makes thousands of identical (clonal) copies of it, and then it synthesizes and secretes antibodies with the same antigen recognition pattern as the BCRs. The activation of B cells corresponding to one specific BCR variant and the dramatic proliferation of that variant is known as clonal selection. This phenomenon drastically, but briefly, changes the proportions of BCR variants expressed by the immune system, and shifts the balance toward BCRs specific to the infecting pathogen.
T and B cells differ in one fundamental way: whereas T cells bind antigens that have been digested and embedded in MHC molecules by APCs, B cells function as APCs that bind intact antigens that have not been processed. Although T and B cells both react with molecules that are termed “antigens,” these lymphocytes actually respond to very different types of molecules. B cells must be able to bind intact antigens because they secrete antibodies that must recognize the pathogen directly, rather than digested remnants of the pathogen. Bacterial carbohydrate and lipid molecules can activate B cells independently from the T cells.
Cytotoxic T Lymphocytes
CTLs, a subclass of T cells, function to clear infections directly. The cell-mediated part of the adaptive immune system consists of CTLs that attack and destroy infected cells. CTLs are particularly important in protecting against viral infections; this is because viruses replicate within cells where they are shielded from extracellular contact with circulating antibodies. When APCs phagocytize pathogens and present MHC I-embedded antigens to naïve CD8+ T cells that express complementary TCRs, the CD8+ T cells become activated to proliferate according to clonal selection. These resulting CTLs then identify non-APCs displaying the same MHC I-embedded antigens (for example, viral proteins)—for example, the CTLs identify infected host cells.
Intracellularly, infected cells typically die after the infecting pathogen replicates to a sufficient concentration and lyses the cell, as many viruses do. CTLs attempt to identify and destroy infected cells before the pathogen can replicate and escape, thereby halting the progression of intracellular infections. CTLs also support NK lymphocytes to destroy early cancers. Cytokines secreted by the TH1 response that stimulates macrophages also stimulate CTLs and enhance their ability to identify and destroy infected cells and tumors.
CTLs sense MHC I-embedded antigens by directly interacting with infected cells via their TCRs. Binding of TCRs with antigens activates CTLs to release perforin and granzyme, degradative enzymes that will induce apoptosis of the infected cell. Recall that this is a similar destruction mechanism to that used by NK cells. In this process, the CTL does not become infected and is not harmed by the secretion of perforin and granzymes. In fact, the functions of NK cells and CTLs are complementary and maximize the removal of infected cells, as illustrated in Figure \(7\). If the NK cell cannot identify the “missing self” pattern of down-regulated MHC I molecules, then the CTL can identify it by the complex of MHC I with foreign antigens, which signals “altered self.” Similarly, if the CTL cannot detect antigen-embedded MHC I because the receptors are depleted from the cell surface, NK cells will destroy the cell instead. CTLs also emit cytokines, such as interferons, that alter surface protein expression in other infected cells, such that the infected cells can be easily identified and destroyed. Moreover, these interferons can also prevent virally infected cells from releasing virus particles.
Exercise \(2\)
Based on what you know about MHC receptors, why do you think an organ transplanted from an incompatible donor to a recipient will be rejected?
Answer
MHC receptors differ from person to person. Thus, MHC receptors on an incompatible donor are considered “non-self” and are rejected by the immune system.
Plasma cells and CTLs are collectively called effector cells: they represent differentiated versions of their naïve counterparts, and they are involved in bringing about the immune defense of killing pathogens and infected host cells.
Mucosal Surfaces and Immune Tolerance
The innate and adaptive immune responses discussed thus far comprise the systemic immune system (affecting the whole body), which is distinct from the mucosal immune system. Mucosal immunity is formed by mucosa-associated lymphoid tissue, which functions independently of the systemic immune system, and which has its own innate and adaptive components. Mucosa-associated lymphoid tissue (MALT), illustrated in Figure \(8\), is a collection of lymphatic tissue that combines with epithelial tissue lining the mucosa throughout the body. This tissue functions as the immune barrier and response in areas of the body with direct contact to the external environment. The systemic and mucosal immune systems use many of the same cell types. Foreign particles that make their way to MALT are taken up by absorptive epithelial cells called M cells and delivered to APCs located directly below the mucosal tissue. M cells function in the transport described, and are located in the Peyer’s patch, a lymphoid nodule. APCs of the mucosal immune system are primarily dendritic cells, with B cells and macrophages having minor roles. Processed antigens displayed on APCs are detected by T cells in the MALT and at various mucosal induction sites, such as the tonsils, adenoids, appendix, or the mesenteric lymph nodes of the intestine. Activated T cells then migrate through the lymphatic system and into the circulatory system to mucosal sites of infection.
MALT is a crucial component of a functional immune system because mucosal surfaces, such as the nasal passages, are the first tissues onto which inhaled or ingested pathogens are deposited. The mucosal tissue includes the mouth, pharynx, and esophagus, and the gastrointestinal, respiratory, and urogenital tracts.
The immune system has to be regulated to prevent wasteful, unnecessary responses to harmless substances, and more importantly so that it does not attack “self.” The acquired ability to prevent an unnecessary or harmful immune response to a detected foreign substance known not to cause disease is described as immune tolerance. Immune tolerance is crucial for maintaining mucosal homeostasis given the tremendous number of foreign substances (such as food proteins) that APCs of the oral cavity, pharynx, and gastrointestinal mucosa encounter. Immune tolerance is brought about by specialized APCs in the liver, lymph nodes, small intestine, and lung that present harmless antigens to an exceptionally diverse population of regulatory T (Treg) cells, specialized lymphocytes that suppress local inflammation and inhibit the secretion of stimulatory immune factors. The combined result of Treg cells is to prevent immunologic activation and inflammation in undesired tissue compartments and to allow the immune system to focus on pathogens instead. In addition to promoting immune tolerance of harmless antigens, other subsets of Treg cells are involved in the prevention of the autoimmune response, which is an inappropriate immune response to host cells or self-antigens. Another Treg class suppresses immune responses to harmful pathogens after the infection has cleared to minimize host cell damage induced by inflammation and cell lysis.
Immunological Memory
The adaptive immune system possesses a memory component that allows for an efficient and dramatic response upon reinvasion of the same pathogen. Memory is handled by the adaptive immune system with little reliance on cues from the innate response. During the adaptive immune response to a pathogen that has not been encountered before, called a primary response, plasma cells secreting antibodies and differentiated T cells increase, then plateau over time. As B and T cells mature into effector cells, a subset of the naïve populations differentiates into B and T memory cells with the same antigen specificities, as illustrated in Figure \(9\).
A memory cell is an antigen-specific B or T lymphocyte that does not differentiate into effector cells during the primary immune response, but that can immediately become effector cells upon re-exposure to the same pathogen. During the primary immune response, memory cells do not respond to antigens and do not contribute to host defenses. As the infection is cleared and pathogenic stimuli subside, the effectors are no longer needed, and they undergo apoptosis. In contrast, the memory cells persist in the circulation.
Exercise \(3\)
The Rh antigen is found on Rh-positive red blood cells. An Rh-negative female can usually carry an Rh-positive fetus to term without difficulty. However, if she has a second Rh-positive fetus, her body may launch an immune attack that causes hemolytic disease of the newborn. Why do you think hemolytic disease is only a problem during the second or subsequent pregnancies?
Answer
If the blood of the mother and fetus mixes, memory cells that recognize the Rh antigen can form late in the first pregnancy. During subsequent pregnancies, these memory cells launch an immune attack on the fetal blood cells. Injection of anti-Rh antibody during the first pregnancy prevents the immune response from occurring.
If the pathogen is never encountered again during the individual’s lifetime, B and T memory cells will circulate for a few years or even several decades and will gradually die off, having never functioned as effector cells. However, if the host is re-exposed to the same pathogen type, circulating memory cells will immediately differentiate into plasma cells and CTLs without input from APCs or TH cells. One reason the adaptive immune response is delayed is because it takes time for naïve B and T cells with the appropriate antigen specificities to be identified and activated. Upon reinfection, this step is skipped, and the result is a more rapid production of immune defenses. Memory B cells that differentiate into plasma cells output tens to hundreds-fold greater antibody amounts than were secreted during the primary response, as the graph in Figure \(10\) illustrates. This rapid and dramatic antibody response may stop the infection before it can even become established, and the individual may not realize they had been exposed.
Vaccination is based on the knowledge that exposure to noninfectious antigens, derived from known pathogens, generates a mild primary immune response. The immune response to vaccination may not be perceived by the host as illness but still confers immune memory. When exposed to the corresponding pathogen to which an individual was vaccinated, the reaction is similar to a secondary exposure. Because each reinfection generates more memory cells and increased resistance to the pathogen, and because some memory cells die, certain vaccine courses involve one or more booster vaccinations to mimic repeat exposures: for instance, tetanus boosters are necessary every ten years because the memory cells only live that long.
Mucosal Immune Memory
A subset of T and B cells of the mucosal immune system differentiates into memory cells just as in the systemic immune system. Upon reinvasion of the same pathogen type, a pronounced immune response occurs at the mucosal site where the original pathogen deposited, but a collective defense is also organized within interconnected or adjacent mucosal tissue. For instance, the immune memory of an infection in the oral cavity would also elicit a response in the pharynx if the oral cavity was exposed to the same pathogen.
Career Connection: Vaccinologist
Vaccination (or immunization) involves the delivery, usually by injection as shown in Figure \(11\), of noninfectious antigen(s) derived from known pathogens. Other components, called adjuvants, are delivered in parallel to help stimulate the immune response. Immunological memory is the reason vaccines work. Ideally, the effect of vaccination is to elicit immunological memory, and thus resistance to specific pathogens without the individual having to experience an infection.
Vaccinologists are involved in the process of vaccine development from the initial idea to the availability of the completed vaccine. This process can take decades, can cost millions of dollars, and can involve many obstacles along the way. For instance, injected vaccines stimulate the systemic immune system, eliciting humoral and cell-mediated immunity, but have little effect on the mucosal response, which presents a challenge because many pathogens are deposited and replicate in mucosal compartments, and the injection does not provide the most efficient immune memory for these disease agents. For this reason, vaccinologists are actively involved in developing new vaccines that are applied via intranasal, aerosol, oral, or transcutaneous (absorbed through the skin) delivery methods. Importantly, mucosal-administered vaccines elicit both mucosal and systemic immunity and produce the same level of disease resistance as injected vaccines.
Currently, a version of intranasal influenza vaccine is available, and the polio and typhoid vaccines can be administered orally, as shown in Figure \(12\). Similarly, the measles and rubella vaccines are being adapted to aerosol delivery using inhalation devices. Eventually, transgenic plants may be engineered to produce vaccine antigens that can be eaten to confer disease resistance. Other vaccines may be adapted to rectal or vaginal application to elicit immune responses in rectal, genitourinary, or reproductive mucosa. Finally, vaccine antigens may be adapted to transdermal application in which the skin is lightly scraped and microneedles are used to pierce the outermost layer. In addition to mobilizing the mucosal immune response, this new generation of vaccines may end the anxiety associated with injections and, in turn, improve patient participation.
Primary Centers of the Immune System
Although the immune system is characterized by circulating cells throughout the body, the regulation, maturation, and intercommunication of immune factors occur at specific sites. The blood circulates immune cells, proteins, and other factors through the body. Approximately 0.1 percent of all cells in the blood are leukocytes, which encompass monocytes (the precursor of macrophages) and lymphocytes. The majority of cells in the blood are erythrocytes (red blood cells). Lymph is a watery fluid that bathes tissues and organs with protective white blood cells and does not contain erythrocytes. Cells of the immune system can travel between the distinct lymphatic and blood circulatory systems, which are separated by interstitial space, by a process called extravasation (passing through to surrounding tissue).
The cells of the immune system originate from hematopoietic stem cells in the bone marrow. Cytokines stimulate these stem cells to differentiate into immune cells. B cell maturation occurs in the bone marrow, whereas naïve T cells transit from the bone marrow to the thymus for maturation. In the thymus, immature T cells that express TCRs complementary to self-antigens are destroyed. This process helps prevent autoimmune responses.
On maturation, T and B lymphocytes circulate to various destinations. Lymph nodes scattered throughout the body, as illustrated in Figure \(13\), house large populations of T and B cells, dendritic cells, and macrophages. Lymph gathers antigens as it drains from tissues. These antigens then are filtered through lymph nodes before the lymph is returned to circulation. APCs in the lymph nodes capture and process antigens and inform nearby lymphocytes about potential pathogens.
The spleen houses B and T cells, macrophages, dendritic cells, and NK cells. The spleen, shown in Figure \(14\), is the site where APCs that have trapped foreign particles in the blood can communicate with lymphocytes. Antibodies are synthesized and secreted by activated plasma cells in the spleen, and the spleen filters foreign substances and antibody-complexed pathogens from the blood. Functionally, the spleen is to the blood as lymph nodes are to the lymph.
Summary
The adaptive immune response is a slower-acting, longer-lasting, and more specific response than the innate response. However, the adaptive response requires information from the innate immune system to function. APCs display antigens via MHC molecules to complementary naïve T cells. In response, the T cells differentiate and proliferate, becoming TH cells or CTLs. TH cells stimulate B cells that have engulfed and presented pathogen-derived antigens. B cells differentiate into plasma cells that secrete antibodies, whereas CTLs induce apoptosis in intracellularly infected or cancerous cells. Memory cells persist after a primary exposure to a pathogen. If re-exposure occurs, memory cells differentiate into effector cells without input from the innate immune system. The mucosal immune system is largely independent from the systemic immune system but functions in a parallel fashion to protect the extensive mucosal surfaces of the body.
Glossary
adaptive immunity
immunity that has memory and occurs after exposure to an antigen either from a pathogen or a vaccination
antigen
foreign or “non-self” protein that triggers the immune response
antigen-presenting cell (APC)
immune cell that detects, engulfs, and informs the adaptive immune response about an infection by presenting the processed antigen on the cell surface
autoimmune response
inappropriate immune response to host cells or self-antigens
cell-mediated immune response
adaptive immune response that is carried out by T cells
clonal selection
activation of B cells corresponding to one specific BCR variant and the dramatic proliferation of that variant
cytotoxic T lymphocyte (CTL)
adaptive immune cell that directly kills infected cells via perforin and granzymes, and releases cytokines to enhance the immune response
dendritic cell
immune cell that processes antigen material and presents it on the surface of other cells to induce an immune response
effector cell
lymphocyte that has differentiated, such as a B cell, plasma cell, or cytotoxic T lymphocyte
epitope
small component of an antigen that is specifically recognized by antibodies, B cells, and T cells; the antigenic determinant
helper T lymphocyte (TH)
cell of the adaptive immune system that binds APCs via MHC II molecules and stimulates B cells or secretes cytokines to initiate the immune response
humoral immune response
adaptive immune response that is controlled by activated B cells and antibodies
immune tolerance
acquired ability to prevent an unnecessary or harmful immune response to a detected foreign body known not to cause disease or to self-antigens
lymph
watery fluid that bathes tissues and organs with protective white blood cells and does not contain erythrocytes
mucosa-associated lymphoid tissue (MALT)
collection of lymphatic tissue that combines with epithelial tissue lining the mucosa throughout the body
memory cell
antigen-specific B or T lymphocyte that does not differentiate into effector cells during the primary immune response but that can immediately become an effector cell upon re-exposure to the same pathogen
plasma cell
immune cell that secrets antibodies; these cells arise from B cells that were stimulated by antigens
regulatory T (Treg) cell
specialized lymphocyte that suppresses local inflammation and inhibits the secretion of cytokines, antibodies, and other stimulatory immune factors; involved in immune tolerance | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/50%3A_The_Immune_System/50.03%3A_Cell-Mediated_Immunity.txt |
Skills to Develop
• Explain cross-reactivity
• Describe the structure and function of antibodies
• Discuss antibody production
An antibody, also known as an immunoglobulin (Ig), is a protein that is produced by plasma cells after stimulation by an antigen. Antibodies are the functional basis of humoral immunity. Antibodies occur in the blood, in gastric and mucus secretions, and in breast milk. Antibodies in these bodily fluids can bind pathogens and mark them for destruction by phagocytes before they can infect cells.
Antibody Structure
An antibody molecule is comprised of four polypeptides: two identical heavy chains (large peptide units) that are partially bound to each other in a “Y” formation, which are flanked by two identical light chains (small peptide units), as illustrated in Figure \(1\). Bonds between the cysteine amino acids in the antibody molecule attach the polypeptides to each other. The areas where the antigen is recognized on the antibody are variable domains and the antibody base is composed of constant domains.
In germ-line B cells, the variable region of the light chain gene has 40 variable (V) and five joining (J) segments. An enzyme called DNA recombinase randomly excises most of these segments out of the gene, and splices one V segment to one J segment. During RNA processing, all but one V and J segment are spliced out. Recombination and splicing may result in over 106 possible VJ combinations. As a result, each differentiated B cell in the human body typically has a unique variable chain. The constant domain, which does not bind antibody, is the same for all antibodies.
Similar to TCRs and BCRs, antibody diversity is produced by the mutation and recombination of approximately 300 different gene segments encoding the light and heavy chain variable domains in precursor cells that are destined to become B cells. The variable domains from the heavy and light chains interact to form the binding site through which an antibody can bind a specific epitope on an antigen. The numbers of repeated constant domains in Ig classes are the same for all antibodies corresponding to a specific class. Antibodies are structurally similar to the extracellular component of the BCRs, and B cell maturation to plasma cells can be visualized in simple terms as the cell acquires the ability to secrete the extracellular portion of its BCR in large quantities.
Antibody Classes
Antibodies can be divided into five classes—IgM, IgG, IgA, IgD, IgE—based on their physiochemical, structural, and immunological properties. IgGs, which make up about 80 percent of all antibodies, have heavy chains that consist of one variable domain and three identical constant domains. IgA and IgD also have three constant domains per heavy chain, whereas IgM and IgE each have four constant domains per heavy chain. The variable domain determines binding specificity and the constant domain of the heavy chain determines the immunological mechanism of action of the corresponding antibody class. It is possible for two antibodies to have the same binding specificities but be in different classes and, therefore, to be involved in different functions.
After an adaptive defense is produced against a pathogen, typically plasma cells first secrete IgM into the blood. BCRs on naïve B cells are of the IgM class and occasionally IgD class. IgM molecules make up approximately ten percent of all antibodies. Prior to antibody secretion, plasma cells assemble IgM molecules into pentamers (five individual antibodies) linked by a joining (J) chain, as shown in Figure \(1\). The pentamer arrangement means that these macromolecules can bind ten identical antigens. However, IgM molecules released early in the adaptive immune response do not bind to antigens as stably as IgGs, which are one of the possible types of antibodies secreted in large quantities upon re-exposure to the same pathogen. Figure \(2\) summarizes the properties of immunoglobulins and illustrates their basic structures.
IgAs populate the saliva, tears, breast milk, and mucus secretions of the gastrointestinal, respiratory, and genitourinary tracts. Collectively, these bodily fluids coat and protect the extensive mucosa (4000 square feet in humans). The total number of IgA molecules in these bodily secretions is greater than the number of IgG molecules in the blood serum. A small amount of IgA is also secreted into the serum in monomeric form. Conversely, some IgM is secreted into bodily fluids of the mucosa. Similar to IgM, IgA molecules are secreted as polymeric structures linked with a J chain. However, IgAs are secreted mostly as dimeric molecules, not pentamers.
IgE is present in the serum in small quantities and is best characterized in its role as an allergy mediator. IgD is also present in small quantities. Similar to IgM, BCRs of the IgD class are found on the surface of naïve B cells. This class supports antigen recognition and maturation of B cells to plasma cells.
Antibody Functions
Differentiated plasma cells are crucial players in the humoral response, and the antibodies they secrete are particularly significant against extracellular pathogens and toxins. Antibodies circulate freely and act independently of plasma cells. Antibodies can be transferred from one individual to another to temporarily protect against infectious disease. For instance, a person who has recently produced a successful immune response against a particular disease agent can donate blood to a nonimmune recipient and confer temporary immunity through antibodies in the donor’s blood serum. This phenomenon is called passive immunity; it also occurs naturally during breastfeeding, which makes breastfed infants highly resistant to infections during the first few months of life.
Antibodies coat extracellular pathogens and neutralize them, as illustrated in Figure \(3\), by blocking key sites on the pathogen that enhance their infectivity (such as receptors that “dock” pathogens on host cells). Antibody neutralization can prevent pathogens from entering and infecting host cells, as opposed to the CTL-mediated approach of killing cells that are already infected to prevent progression of an established infection. The neutralized antibody-coated pathogens can then be filtered by the spleen and eliminated in urine or feces.
Antibodies also mark pathogens for destruction by phagocytic cells, such as macrophages or neutrophils, because phagocytic cells are highly attracted to macromolecules complexed with antibodies. Phagocytic enhancement by antibodies is called opsonization. In a process called complement fixation, IgM and IgG in serum bind to antigens and provide docking sites onto which sequential complement proteins can bind. The combination of antibodies and complement enhances opsonization even further and promotes rapid clearing of pathogens.
Affinity, Avidity, and Cross Reactivity
Not all antibodies bind with the same strength, specificity, and stability. In fact, antibodies exhibit different affinities (attraction) depending on the molecular complementarity between antigen and antibody molecules, as illustrated in Figure \(4\). An antibody with a higher affinity for a particular antigen would bind more strongly and stably, and thus would be expected to present a more challenging defense against the pathogen corresponding to the specific antigen.
The term avidity describes binding by antibody classes that are secreted as joined, multivalent structures (such as IgM and IgA). Although avidity measures the strength of binding, just as affinity does, the avidity is not simply the sum of the affinities of the antibodies in a multimeric structure. The avidity depends on the number of identical binding sites on the antigen being detected, as well as other physical and chemical factors. Typically, multimeric antibodies, such as pentameric IgM, are classified as having lower affinity than monomeric antibodies, but high avidity. Essentially, the fact that multimeric antibodies can bind many antigens simultaneously balances their slightly lower binding strength for each antibody/antigen interaction.
Antibodies secreted after binding to one epitope on an antigen may exhibit cross reactivity for the same or similar epitopes on different antigens. Because an epitope corresponds to such a small region (the surface area of about four to six amino acids), it is possible for different macromolecules to exhibit the same molecular identities and orientations over short regions. Cross reactivity describes when an antibody binds not to the antigen that elicited its synthesis and secretion, but to a different antigen.
Cross reactivity can be beneficial if an individual develops immunity to several related pathogens despite having only been exposed to or vaccinated against one of them. For instance, antibody cross reactivity may occur against the similar surface structures of various Gram-negative bacteria. Conversely, antibodies raised against pathogenic molecular components that resemble self molecules may incorrectly mark host cells for destruction and cause autoimmune damage. Patients who develop systemic lupus erythematosus (SLE) commonly exhibit antibodies that react with their own DNA. These antibodies may have been initially raised against the nucleic acid of microorganisms but later cross-reacted with self-antigens. This phenomenon is also called molecular mimicry.
Antibodies of the Mucosal Immune System
Antibodies synthesized by the mucosal immune system include IgA and IgM. Activated B cells differentiate into mucosal plasma cells that synthesize and secrete dimeric IgA, and to a lesser extent, pentameric IgM. Secreted IgA is abundant in tears, saliva, breast milk, and in secretions of the gastrointestinal and respiratory tracts. Antibody secretion results in a local humoral response at epithelial surfaces and prevents infection of the mucosa by binding and neutralizing pathogens.
Summary
Antibodies (immunoglobulins) are the molecules secreted from plasma cells that mediate the humoral immune response. There are five antibody classes; an antibody's class determines its mechanism of action and production site but does not control its binding specificity. Antibodies bind antigens via variable domains and can either neutralize pathogens or mark them for phagocytosis or activate the complement cascade.
Glossary
affinity
attraction of molecular complementarity between antigen and antibody molecules
antibody
protein that is produced by plasma cells after stimulation by an antigen; also known as an immunoglobulin
avidity
total binding strength of a multivalent antibody with antigen
cross reactivity
binding of an antibody to an epitope corresponding to an antigen that is different from the one the antibody was raised against
passive immunity
transfer of antibodies from one individual to another to provide temporary protection against pathogens | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/50%3A_The_Immune_System/50.04%3A_Humoral_Immunity_and_Antibody_Production.txt |
Skills to Develop
• Describe hypersensitivity
• Define autoimmunity
A functioning immune system is essential for survival, but even the sophisticated cellular and molecular defenses of the mammalian immune response can be defeated by pathogens at virtually every step. In the competition between immune protection and pathogen evasion, pathogens have the advantage of more rapid evolution because of their shorter generation time and other characteristics. For instance, Streptococcus pneumoniae (bacterium that cause pneumonia and meningitis) surrounds itself with a capsule that inhibits phagocytes from engulfing it and displaying antigens to the adaptive immune system. Staphylococcus aureus (bacterium that can cause skin infections, abscesses, and meningitis) synthesizes a toxin called leukocidin that kills phagocytes after they engulf the bacterium. Other pathogens can also hinder the adaptive immune system. HIV infects TH cells via their CD4 surface molecules, gradually depleting the number of TH cells in the body; this inhibits the adaptive immune system’s capacity to generate sufficient responses to infection or tumors. As a result, HIV-infected individuals often suffer from infections that would not cause illness in people with healthy immune systems but which can cause devastating illness to immune-compromised individuals. Maladaptive responses of immune cells and molecules themselves can also disrupt the proper functioning of the entire system, leading to host cell damage that could become fatal.
Immunodeficiency
Failures, insufficiencies, or delays at any level of the immune response can allow pathogens or tumor cells to gain a foothold and replicate or proliferate to high enough levels that the immune system becomes overwhelmed. Immunodeficiency is the failure, insufficiency, or delay in the response of the immune system, which may be acquired or inherited. Immunodeficiency can be acquired as a result of infection with certain pathogens (such as HIV), chemical exposure (including certain medical treatments), malnutrition, or possibly by extreme stress. For instance, radiation exposure can destroy populations of lymphocytes and elevate an individual’s susceptibility to infections and cancer. Dozens of genetic disorders result in immunodeficiencies, including Severe Combined Immunodeficiency (SCID), Bare lymphocyte syndrome, and MHC II deficiencies. Rarely, primary immunodeficiencies that are present from birth may occur. Neutropenia is one form in which the immune system produces a below-average number of neutrophils, the body’s most abundant phagocytes. As a result, bacterial infections may go unrestricted in the blood, causing serious complications.
Hypersensitivities
Maladaptive immune responses toward harmless foreign substances or self antigens that occur after tissue sensitization are termed hypersensitivities. The types of hypersensitivities include immediate, delayed, and autoimmunity. A large proportion of the population is affected by one or more types of hypersensitivity.
Allergies
The immune reaction that results from immediate hypersensitivities in which an antibody-mediated immune response occurs within minutes of exposure to a harmless antigen is called an allergy. In the United States, 20 percent of the population exhibits symptoms of allergy or asthma, whereas 55 percent test positive against one or more allergens. Upon initial exposure to a potential allergen, an allergic individual synthesizes antibodies of the IgE class via the typical process of APCs presenting processed antigen to TH cells that stimulate B cells to produce IgE. This class of antibodies also mediates the immune response to parasitic worms. The constant domain of the IgE molecules interact with mast cells embedded in connective tissues. This process primes, or sensitizes, the tissue. Upon subsequent exposure to the same allergen, IgE molecules on mast cells bind the antigen via their variable domains and stimulate the mast cell to release the modified amino acids histamine and serotonin; these chemical mediators then recruit eosinophils which mediate allergic responses. Figure \(1\) shows an example of an allergic response to ragweed pollen. The effects of an allergic reaction range from mild symptoms like sneezing and itchy, watery eyes to more severe or even life-threatening reactions involving intensely itchy welts or hives, airway contraction with severe respiratory distress, and plummeting blood pressure. This extreme reaction is known as anaphylactic shock. If not treated with epinephrine to counter the blood pressure and breathing effects, this condition can be fatal.
Delayed hypersensitivity is a cell-mediated immune response that takes approximately one to two days after secondary exposure for a maximal reaction to be observed. This type of hypersensitivity involves the TH1 cytokine-mediated inflammatory response and may manifest as local tissue lesions or contact dermatitis (rash or skin irritation). Delayed hypersensitivity occurs in some individuals in response to contact with certain types of jewelry or cosmetics. Delayed hypersensitivity facilitates the immune response to poison ivy and is also the reason why the skin test for tuberculosis results in a small region of inflammation on individuals who were previously exposed to Mycobacterium tuberculosis. That is also why cortisone is used to treat such responses: it will inhibit cytokine production.
Autoimmunity
Autoimmunity is a type of hypersensitivity to self antigens that affects approximately five percent of the population. Most types of autoimmunity involve the humoral immune response. Antibodies that inappropriately mark self components as foreign are termed autoantibodies. In patients with the autoimmune disease myasthenia gravis, muscle cell receptors that induce contraction in response to acetylcholine are targeted by antibodies. The result is muscle weakness that may include marked difficultly with fine and/or gross motor functions. In systemic lupus erythematosus, a diffuse autoantibody response to the individual’s own DNA and proteins results in various systemic diseases. As illustrated in Figure \(2\), systemic lupus erythematosus may affect the heart, joints, lungs, skin, kidneys, central nervous system, or other tissues, causing tissue damage via antibody binding, complement recruitment, lysis, and inflammation.
Autoimmunity can develop with time, and its causes may be rooted in molecular mimicry. Antibodies and TCRs may bind self antigens that are structurally similar to pathogen antigens, which the immune receptors first raised. As an example, infection with Streptococcus pyogenes (bacterium that causes strep throat) may generate antibodies or T cells that react with heart muscle, which has a similar structure to the surface of S. pyogenes. These antibodies can damage heart muscle with autoimmune attacks, leading to rheumatic fever. Insulin-dependent (Type 1) diabetes mellitus arises from a destructive inflammatory TH1 response against insulin-producing cells of the pancreas. Patients with this autoimmunity must be injected with insulin that originates from other sources.
Summary
Immune disruptions may involve insufficient immune responses or inappropriate immune targets. Immunodeficiency increases an individual's susceptibility to infections and cancers. Hypersensitivities are misdirected responses either to harmless foreign particles, as in the case of allergies, or to host factors, as in the case of autoimmunity. Reactions to self components may be the result of molecular mimicry.
Glossary
allergy
immune reaction that results from immediate hypersensitivities in which an antibody-mediated immune response occurs within minutes of exposure to a harmless antigen
autoantibody
antibody that incorrectly marks “self” components as foreign and stimulates the immune response
autoimmunity
type of hypersensitivity to self antigens
hypersensitivities
spectrum of maladaptive immune responses toward harmless foreign particles or self antigens; occurs after tissue sensitization and includes immediate-type (allergy), delayed-type, and autoimmunity
immunodeficiency
failure, insufficiency, or delay at any level of the immune system, which may be acquired or inherited | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/50%3A_The_Immune_System/50.05%3A_Autoimmunity_and_Hypersensitivity.txt |
Skills to Develop
• Identify major viral illnesses that affect humans
• Compare vaccinations and anti-viral drugs as medical approaches to viruses
Viruses cause a variety of diseases in animals, including humans, ranging from the common cold to potentially fatal illnesses like meningitis (Figure \(1\)). These diseases can be treated by antiviral drugs or by vaccines, but some viruses, such as HIV, are capable of both avoiding the immune response and mutating to become resistant to antiviral drugs.
Vaccines for Prevention
While we do have limited numbers of effective antiviral drugs, such as those used to treat HIV and influenza, the primary method of controlling viral disease is by vaccination, which is intended to prevent outbreaks by building immunity to a virus or virus family (Figure\(1\)). Vaccines may be prepared using live viruses, killed viruses, or molecular subunits of the virus. The killed viral vaccines and subunit viruses are both incapable of causing disease.
Live viral vaccines are designed in the laboratory to cause few symptoms in recipients while giving them protective immunity against future infections. Polio was one disease that represented a milestone in the use of vaccines. Mass immunization campaigns in the 1950s (killed vaccine) and 1960s (live vaccine) significantly reduced the incidence of the disease, which caused muscle paralysis in children and generated a great amount of fear in the general population when regional epidemics occurred. The success of the polio vaccine paved the way for the routine dispensation of childhood vaccines against measles, mumps, rubella, chickenpox, and other diseases.
The danger of using live vaccines, which are usually more effective than killed vaccines, is the low but significant danger that these viruses will revert to their disease-causing form by back mutations. Live vaccines are usually made by attenuating (weakening) the “wild-type” (disease-causing) virus by growing it in the laboratory in tissues or at temperatures different from what the virus is accustomed to in the host. Adaptations to these new cells or temperatures induce mutations in the genomes of the virus, allowing it to grow better in the laboratory while inhibiting its ability to cause disease when reintroduced into conditions found in the host. These attenuated viruses thus still cause infection, but they do not grow very well, allowing the immune response to develop in time to prevent major disease. Back mutations occur when the vaccine undergoes mutations in the host such that it readapts to the host and can again cause disease, which can then be spread to other humans in an epidemic. This type of scenario happened as recently as 2007 in Nigeria where mutations in a polio vaccine led to an epidemic of polio in that country.
Some vaccines are in continuous development because certain viruses, such as influenza and HIV, have a high mutation rate compared to other viruses and normal host cells. With influenza, mutations in the surface molecules of the virus help the organism evade the protective immunity that may have been obtained in a previous influenza season, making it necessary for individuals to get vaccinated every year. Other viruses, such as those that cause the childhood diseases measles, mumps, and rubella, mutate so infrequently that the same vaccine is used year after year.
Link to Learning
Watch this NOVA video to learn how microbiologists are attempting to replicate the deadly 1918 Spanish influenza virus so they can understand more about virology.
Vaccines and Anti-viral Drugs for Treatment
In some cases, vaccines can be used to treat an active viral infection. The concept behind this is that by giving the vaccine, immunity is boosted without adding more disease-causing virus. In the case of rabies, a fatal neurological disease transmitted via the saliva of rabies virus-infected animals, the progression of the disease from the time of the animal bite to the time it enters the central nervous system may be 2 weeks or longer. This is enough time to vaccinate an individual who suspects that they have been bitten by a rabid animal, and their boosted immune response is sufficient to prevent the virus from entering nervous tissue. Thus, the potentially fatal neurological consequences of the disease are averted, and the individual only has to recover from the infected bite. This approach is also being used for the treatment of Ebola, one of the fastest and most deadly viruses on earth. Transmitted by bats and great apes, this disease can cause death in 70–90 percent of infected humans within 2 weeks. Using newly developed vaccines that boost the immune response in this way, there is hope that affected individuals will be better able to control the virus, potentially saving a greater percentage of infected persons from a rapid and very painful death.
Another way of treating viral infections is the use of antiviral drugs. These drugs often have limited success in curing viral disease, but in many cases, they have been used to control and reduce symptoms for a wide variety of viral diseases. For most viruses, these drugs can inhibit the virus by blocking the actions of one or more of its proteins. It is important that the targeted proteins be encoded by viral genes and that these molecules are not present in a healthy host cell. In this way, viral growth is inhibited without damaging the host. There are large numbers of antiviral drugs available to treat infections, some specific for a particular virus and others that can affect multiple viruses.
Antivirals have been developed to treat genital herpes (herpes simplex II) and influenza. For genital herpes, drugs such as acyclovir can reduce the number and duration of episodes of active viral disease, during which patients develop viral lesions in their skin cells. As the virus remains latent in nervous tissue of the body for life, this drug is not curative but can make the symptoms of the disease more manageable. For influenza, drugs like Tamiflu (oseltamivir) (Figure \(3\)) can reduce the duration of “flu” symptoms by 1 or 2 days, but the drug does not prevent symptoms entirely. Tamiflu works by inhibiting an enzyme (viral neuraminidase) that allows new virions to leave their infected cells. Thus, Tamiflu inhibits the spread of virus from infected to uninfected cells. Other antiviral drugs, such as Ribavirin, have been used to treat a variety of viral infections, although its mechanism of action against certain viruses remains unclear.
By far, the most successful use of antivirals has been in the treatment of the retrovirus HIV, which causes a disease that, if untreated, is usually fatal within 10–12 years after infection. Anti-HIV drugs have been able to control viral replication to the point that individuals receiving these drugs survive for a significantly longer time than the untreated.
Anti-HIV drugs inhibit viral replication at many different phases of the HIV replicative cycle (Figure \(4\)). Drugs have been developed that inhibit the fusion of the HIV viral envelope with the plasma membrane of the host cell (fusion inhibitors), the conversion of its RNA genome into double-stranded DNA (reverse transcriptase inhibitors), the integration of the viral DNA into the host genome (integrase inhibitors), and the processing of viral proteins (protease inhibitors).
When any of these drugs are used individually, the high mutation rate of the virus allows it to easily and rapidly develop resistance to the drug, limiting the drug’s effectiveness. The breakthrough in the treatment of HIV was the development of HAART, highly active anti-retroviral therapy, which involves a mixture of different drugs, sometimes called a drug “cocktail.” By attacking the virus at different stages of its replicative cycle, it is much more difficult for the virus to develop resistance to multiple drugs at the same time. Still, even with the use of combination HAART therapy, there is concern that, over time, the virus will develop resistance to this therapy. Thus, new anti-HIV drugs are constantly being developed with the hope of continuing the battle against this highly fatal virus.
Everyday Connection: Applied Virology
The study of viruses has led to the development of a variety of new ways to treat non-viral diseases. Viruses have been used in gene therapy. Gene therapy is used to treat genetic diseases such as severe combined immunodeficiency (SCID), a heritable, recessive disease in which children are born with severely compromised immune systems. One common type of SCID is due to the lack of an enzyme, adenosine deaminase (ADA), which breaks down purine bases. To treat this disease by gene therapy, bone marrow cells are taken from a SCID patient and the ADA gene is inserted. This is where viruses come in, and their use relies on their ability to penetrate living cells and bring genes in with them. Viruses such as adenovirus, an upper respiratory human virus, are modified by the addition of the ADA gene, and the virus then transports this gene into the cell. The modified cells, now capable of making ADA, are then given back to the patients in the hope of curing them. Gene therapy using viruses as carrier of genes (viral vectors), although still experimental, holds promise for the treatment of many genetic diseases. Still, many technological problems need to be solved for this approach to be a viable method for treating genetic disease.
Another medical use for viruses relies on their specificity and ability to kill the cells they infect. Oncolytic viruses are engineered in the laboratory specifically to attack and kill cancer cells. A genetically modified adenovirus known as H101 has been used since 2005 in clinical trials in China to treat head and neck cancers. The results have been promising, with a greater short-term response rate to the combination of chemotherapy and viral therapy than to chemotherapy treatment alone. This ongoing research may herald the beginning of a new age of cancer therapy, where viruses are engineered to find and specifically kill cancer cells, regardless of where in the body they may have spread.
A third use of viruses in medicine relies on their specificity and involves using bacteriophages in the treatment of bacterial infections. Bacterial diseases have been treated with antibiotics since the 1940s. However, over time, many bacteria have developed resistance to antibiotics. A good example is methicillin-resistant Staphylococcus aureus (MRSA, pronounced “mersa”), an infection commonly acquired in hospitals. This bacterium is resistant to a variety of antibiotics, making it difficult to treat. The use of bacteriophages specific for such bacteria would bypass their resistance to antibiotics and specifically kill them. Although phage therapy is in use in the Republic of Georgia to treat antibiotic-resistant bacteria, its use to treat human diseases has not been approved in most countries. However, the safety of the treatment was confirmed in the United States when the U.S. Food and Drug Administration approved spraying meats with bacteriophages to destroy the food pathogen Listeria. As more and more antibiotic-resistant strains of bacteria evolve, the use of bacteriophages might be a potential solution to the problem, and the development of phage therapy is of much interest to researchers worldwide.
Summary
Viruses cause a variety of diseases in humans. Many of these diseases can be prevented by the use of viral vaccines, which stimulate protective immunity against the virus without causing major disease. Viral vaccines may also be used in active viral infections, boosting the ability of the immune system to control or destroy the virus. A series of antiviral drugs that target enzymes and other protein products of viral genes have been developed and used with mixed success. Combinations of anti-HIV drugs have been used to effectively control the virus, extending the lifespans of infected individuals. Viruses have many uses in medicines, such as in the treatment of genetic disorders, cancer, and bacterial infections.
Glossary
attenuation
weakening of a virus during vaccine development
back mutation
when a live virus vaccine reverts back to it disease-causing phenotype
gene therapy
treatment of genetic disease by adding genes, using viruses to carry the new genes inside the cell
oncolytic virus
virus engineered to specifically infect and kill cancer cells
phage therapy
treatment of bacterial diseases using bacteriophages specific to a particular bacterium
vaccine
weakened solution of virus components, viruses, or other agents that produce an immune response | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/50%3A_The_Immune_System/50.07%3A_Pathogens_that_Evade_the_Immune_System.txt |
• 51.1: Animal Reproductive Strategies
During sexual reproduction the genetic material of two individuals is combined to produce genetically diverse offspring that differ from their parents. The genetic diversity of sexually produced offspring is thought to give species a better chance of surviving in an unpredictable or changing environment. Species that reproduce sexually must maintain two different types of individuals, males and females, which can limit the ability to colonize new habitats as both sexes must be present.
• 51.2: Vertebrate Fertilization and Development
Sexual reproduction starts with the combination of a sperm and an egg in a process called fertilization. This can occur either inside (internal fertilization) or outside (external fertilization) the body of the female. Humans provide an example of the former whereas seahorse reproduction is an example of the latter.
• 51.3: Structure and Function of the Human Male Reproductive System
As animals became more complex, specific organs and organ systems developed to support specific functions for the organism. The reproductive structures that evolved in land animals allow males and females to mate, fertilize internally, and support the growth and development of offspring.
• 51.4: Structure and Function of the Female Reproductive System
As animals became more complex, specific organs and organ systems developed to support specific functions for the organism. The reproductive structures that evolved in land animals allow males and females to mate, fertilize internally, and support the growth and development of offspring.
• 51.5: Contraception and Fertility Treatments
Pregnancy begins with the fertilization of an egg and continues through to the birth of the individual. The length of time of gestation varies among animals, but is very similar among the great apes: human gestation is 266 days, while chimpanzee gestation is 237 days, a gorilla’s is 257 days, and orangutan gestation is 260 days long. The fox has a 57-day gestation. Dogs and cats have similar gestations averaging 60 days.
51: The Reproductive System
Skills to Develop
• Describe advantages and disadvantages of asexual and sexual reproduction
• Discuss asexual reproduction methods
• Discuss sexual reproduction methods
Animals produce offspring through asexual and/or sexual reproduction. Both methods have advantages and disadvantages. Asexual reproduction produces offspring that are genetically identical to the parent because the offspring are all clones of the original parent. A single individual can produce offspring asexually and large numbers of offspring can be produced quickly. In a stable or predictable environment, asexual reproduction is an effective means of reproduction because all the offspring will be adapted to that environment. In an unstable or unpredictable environment asexually-reproducing species may be at a disadvantage because all the offspring are genetically identical and may not have the genetic variation to survive in new or different conditions. On the other hand, the rapid rates of asexual reproduction may allow for a speedy response to environmental changes if individuals have mutations. An additional advantage of asexual reproduction is that colonization of new habitats may be easier when an individual does not need to find a mate to reproduce.
During sexual reproduction the genetic material of two individuals is combined to produce genetically diverse offspring that differ from their parents. The genetic diversity of sexually produced offspring is thought to give species a better chance of surviving in an unpredictable or changing environment. Species that reproduce sexually must maintain two different types of individuals, males and females, which can limit the ability to colonize new habitats as both sexes must be present.
Asexual Reproduction
Asexual reproduction occurs in prokaryotic microorganisms (bacteria) and in some eukaryotic single-celled and multi-celled organisms. There are a number of ways that animals reproduce asexually.
Fission
Fission, also called binary fission, occurs in prokaryotic microorganisms and in some invertebrate, multi-celled organisms. After a period of growth, an organism splits into two separate organisms. Some unicellular eukaryotic organisms undergo binary fission by mitosis. In other organisms, part of the individual separates and forms a second individual. This process occurs, for example, in many asteroid echinoderms through splitting of the central disk. Some sea anemones and some coral polyps (Figure \(1\)) also reproduce through fission.
Budding
Budding is a form of asexual reproduction that results from the outgrowth of a part of a cell or body region leading to a separation from the original organism into two individuals. Budding occurs commonly in some invertebrate animals such as corals and hydras. In hydras, a bud forms that develops into an adult and breaks away from the main body, as illustrated in Figure \(2\), whereas in coral budding, the bud does not detach and multiplies as part of a new colony.
Link to Learning
Watch a video of a hydra budding.
Fragmentation
Fragmentation is the breaking of the body into two parts with subsequent regeneration. If the animal is capable of fragmentation, and the part is big enough, a separate individual will regrow.
For example, in many sea stars, asexual reproduction is accomplished by fragmentation. Figure \(3\) illustrates a sea star for which an arm of the individual is broken off and regenerates a new sea star. Fisheries workers have been known to try to kill the sea stars eating their clam or oyster beds by cutting them in half and throwing them back into the ocean. Unfortunately for the workers, the two parts can each regenerate a new half, resulting in twice as many sea stars to prey upon the oysters and clams. Fragmentation also occurs in annelid worms, turbellarians, and poriferans.
Note that in fragmentation, there is generally a noticeable difference in the size of the individuals, whereas in fission, two individuals of approximate size are formed.
Parthenogenesis
Parthenogenesis is a form of asexual reproduction where an egg develops into a complete individual without being fertilized. The resulting offspring can be either haploid or diploid, depending on the process and the species. Parthenogenesis occurs in invertebrates such as water flees, rotifers, aphids, stick insects, some ants, wasps, and bees. Bees use parthenogenesis to produce haploid males (drones) and diploid females (workers). If an egg is fertilized, a queen is produced. The queen bee controls the reproduction of the hive bees to regulate the type of bee produced.
Some vertebrate animals—such as certain reptiles, amphibians, and fish—also reproduce through parthenogenesis. Although more common in plants, parthenogenesis has been observed in animal species that were segregated by sex in terrestrial or marine zoos. Two female Komodo dragons, a hammerhead shark, and a blacktop shark have produced parthenogenic young when the females have been isolated from males.
Sexual Reproduction
Sexual reproduction is the combination of (usually haploid) reproductive cells from two individuals to form a third (usually diploid) unique offspring. Sexual reproduction produces offspring with novel combinations of genes. This can be an adaptive advantage in unstable or unpredictable environments. As humans, we are used to thinking of animals as having two separate sexes—male and female—determined at conception. However, in the animal kingdom, there are many variations on this theme.
Hermaphroditism
Hermaphroditism occurs in animals where one individual has both male and female reproductive parts. Invertebrates such as earthworms, slugs, tapeworms and snails, shown in Figure \(4\), are often hermaphroditic. Hermaphrodites may self-fertilize or may mate with another of their species, fertilizing each other and both producing offspring. Self fertilization is common in animals that have limited mobility or are not motile, such as barnacles and clams.
Sex Determination
Mammalian sex determination is determined genetically by the presence of X and Y chromosomes. Individuals homozygous for X (XX) are female and heterozygous individuals (XY) are male. The presence of a Y chromosome causes the development of male characteristics and its absence results in female characteristics. The XY system is also found in some insects and plants.
Avian sex determination is dependent on the presence of Z and W chromosomes. Homozygous for Z (ZZ) results in a male and heterozygous (ZW) results in a female. The W appears to be essential in determining the sex of the individual, similar to the Y chromosome in mammals. Some fish, crustaceans, insects (such as butterflies and moths), and reptiles use this system.
The sex of some species is not determined by genetics but by some aspect of the environment. Sex determination in some crocodiles and turtles, for example, is often dependent on the temperature during critical periods of egg development. This is referred to as environmental sex determination, or more specifically as temperature-dependent sex determination. In many turtles, cooler temperatures during egg incubation produce males and warm temperatures produce females. In some crocodiles, moderate temperatures produce males and both warm and cool temperatures produce females. In some species, sex is both genetic- and temperature-dependent.
Individuals of some species change their sex during their lives, alternating between male and female. If the individual is female first, it is termed protogyny or “first female,” if it is male first, its termed protandry or “first male.” Oysters, for example, are born male, grow, and become female and lay eggs; some oyster species change sex multiple times.
Summary
Reproduction may be asexual when one individual produces genetically identical offspring, or sexual when the genetic material from two individuals is combined to produce genetically diverse offspring. Asexual reproduction occurs through fission, budding, and fragmentation. Sexual reproduction may mean the joining of sperm and eggs within animals’ bodies or it may mean the release of sperm and eggs into the environment. An individual may be one sex, or both; it may start out as one sex and switch during its life, or it may stay male or female.
Glossary
asexual reproduction
form of reproduction that produces offspring that are genetically identical to the parent
budding
form of asexual reproduction that results from the outgrowth of a part of a cell leading to a separation from the original animal into two individuals
fission
(also, binary fission) method by which multicellular organisms increase in size or asexual reproduction in which a unicellular organism splits into two separate organisms by mitosis
fragmentation
cutting or fragmenting of the original animal into parts and the growth of a separate animal from each part
hermaphroditism
state of having both male and female reproductive parts within the same individual
parthenogenesis
form of asexual reproduction where an egg develops into a complete individual without being fertilized
sexual reproduction
mixing of genetic material from two individuals to produce genetically unique offspring | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/51%3A_The_Reproductive_System/51.01%3A_Animal_Reproductive_Strategies.txt |
Skills to Develop
• Discuss internal and external methods of fertilization
• Describe the methods used by animals for development of offspring during gestation
• Describe the anatomical adaptions that occurred in animals to facilitate reproduction
Sexual reproduction starts with the combination of a sperm and an egg in a process called fertilization. This can occur either inside (internal fertilization) or outside (external fertilization) the body of the female. Humans provide an example of the former whereas seahorse reproduction is an example of the latter.
External Fertilization
External fertilization usually occurs in aquatic environments where both eggs and sperm are released into the water. After the sperm reaches the egg, fertilization takes place. Most external fertilization happens during the process of spawning where one or several females release their eggs and the male(s) release sperm in the same area, at the same time. The release of the reproductive material may be triggered by water temperature or the length of daylight. Nearly all fish spawn, as do crustaceans (such as crabs and shrimp), mollusks (such as oysters), squid, and echinoderms (such as sea urchins and sea cucumbers). Figure \(1\) shows salmon spawning in a shallow stream. Frogs, like those shown in Figure \(2\), corals, molluscs, and sea cucumbers also spawn.
Pairs of fish that are not broadcast spawners may exhibit courtship behavior. This allows the female to select a particular male. The trigger for egg and sperm release (spawning) causes the egg and sperm to be placed in a small area, enhancing the possibility of fertilization.
External fertilization in an aquatic environment protects the eggs from drying out. Broadcast spawning can result in a greater mixture of the genes within a group, leading to higher genetic diversity and a greater chance of species survival in a hostile environment. For sessile aquatic organisms like sponges, broadcast spawning is the only mechanism for fertilization and colonization of new environments. The presence of the fertilized eggs and developing young in the water provides opportunities for predation resulting in a loss of offspring. Therefore, millions of eggs must be produced by individuals, and the offspring produced through this method must mature rapidly. The survival rate of eggs produced through broadcast spawning is low.
Internal Fertilization
Internal fertilization occurs most often in land-based animals, although some aquatic animals also use this method. There are three ways that offspring are produced following internal fertilization. In oviparity, fertilized eggs are laid outside the female’s body and develop there, receiving nourishment from the yolk that is a part of the egg. This occurs in most bony fish, many reptiles, some cartilaginous fish, most amphibians, two mammals, and all birds. Reptiles and insects produce leathery eggs, while birds and turtles produce eggs with high concentrations of calcium carbonate in the shell, making them hard. Chicken eggs are an example of this second type.
In ovoviparity, fertilized eggs are retained in the female, but the embryo obtains its nourishment from the egg’s yolk and the young are fully developed when they are hatched. This occurs in some bony fish (like the guppy Lebistes reticulatus), some sharks, some lizards, some snakes (such as the garter snake Thamnophis sirtalis), some vipers, and some invertebrate animals (like the Madagascar hissing cockroach Gromphadorhina portentosa).
In viviparity the young develop within the female, receiving nourishment from the mother’s blood through a placenta. The offspring develops in the female and is born alive. This occurs in most mammals, some cartilaginous fish, and a few reptiles.
Internal fertilization has the advantage of protecting the fertilized egg from dehydration on land. The embryo is isolated within the female, which limits predation on the young. Internal fertilization enhances the fertilization of eggs by a specific male. Fewer offspring are produced through this method, but their survival rate is higher than that for external fertilization.
The Evolution of Reproduction
Once multicellular organisms evolved and developed specialized cells, some also developed tissues and organs with specialized functions. An early development in reproduction occurred in the Annelids. These organisms produce sperm and eggs from undifferentiated cells in their coelom and store them in that cavity. When the coelom becomes filled, the cells are released through an excretory opening or by the body splitting open. Reproductive organs evolved with the development of gonads that produce sperm and eggs. These cells went through meiosis, an adaption of mitosis, which reduced the number of chromosomes in each reproductive cell by half, while increasing the number of cells through cell division.
Complete reproductive systems were developed in insects, with separate sexes. Sperm are made in testes and then travel through coiled tubes to the epididymis for storage. Eggs mature in the ovary. When they are released from the ovary, they travel to the uterine tubes for fertilization. Some insects have a specialized sac, called a spermatheca, which stores sperm for later use, sometimes up to a year. Fertilization can be timed with environmental or food conditions that are optimal for offspring survival.
Vertebrates have similar structures, with a few differences. Non-mammals, such as birds and reptiles, have a common body opening, called a cloaca, for the digestive, excretory and reproductive systems. Coupling between birds usually involves positioning the cloaca openings opposite each other for transfer of sperm. Mammals have separate openings for the systems in the female and a uterus for support of developing offspring. The uterus has two chambers in species that produce large numbers of offspring at a time, while species that produce one offspring, such as primates, have a single uterus.
Sperm transfer from the male to the female during reproduction ranges from releasing the sperm into the watery environment for external fertilization, to the joining of cloaca in birds, to the development of a penis for direct delivery into the female’s vagina in mammals.
Summary
Sexual reproduction starts with the combination of a sperm and an egg in a process called fertilization. This can occur either outside the bodies or inside the female. Both methods have advantages and disadvantages. Once fertilized, the eggs can develop inside the female or outside. If the egg develops outside the body, it usually has a protective covering over it. Animal anatomy evolved various ways to fertilize, hold, or expel the egg. The method of fertilization varies among animals. Some species release the egg and sperm into the environment, some species retain the egg and receive the sperm into the female body and then expel the developing embryo covered with shell, while still other species retain the developing offspring through the gestation period.
Glossary
cloaca
common body opening for the digestive, excretory, and reproductive systems found in non-mammals, such as birds
external fertilization
fertilization of egg by sperm outside animal body, often during spawning
internal fertilization
fertilization of egg by sperm inside the body of the female
oviparity
process by which fertilized eggs are laid outside the female’s body and develop there, receiving nourishment from the yolk that is a part of the egg
ovoviparity
process by which fertilized eggs are retained within the female; the embryo obtains its nourishment from the egg’s yolk and the young are fully developed when they are hatched
spermatheca
specialized sac that stores sperm for later use
viviparity
process in which the young develop within the female, receiving nourishment from the mother’s blood through a placenta | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/51%3A_The_Reproductive_System/51.02%3A_Vertebrate_Fertilization_and_Development.txt |
Skills to Develop
• Describe human male and female reproductive anatomies
• Discuss the human sexual response
• Describe spermatogenesis and oogenesis and discuss their differences and similarities
As animals became more complex, specific organs and organ systems developed to support specific functions for the organism. The reproductive structures that evolved in land animals allow males and females to mate, fertilize internally, and support the growth and development of offspring.
Human Reproductive Anatomy
The reproductive tissues of male and female humans develop similarly in utero until a low level of the hormone testosterone is released from male gonads. Testosterone causes the undeveloped tissues to differentiate into male sexual organs. When testosterone is absent, the tissues develop into female sexual tissues. Primitive gonads become testes or ovaries. Tissues that produce a penis in males produce a clitoris in females. The tissue that will become the scrotum in a male becomes the labia in a female; that is, they are homologous structures.
Male Reproductive Anatomy
In the male reproductive system, the scrotum houses the testicles or testes (singular: testis), including providing passage for blood vessels, nerves, and muscles related to testicular function. The testes are a pair of male reproductive organs that produce sperm and some reproductive hormones. Each testis is approximately 2.5 by 3.8 cm (1.5 by 1 in) in size and divided into wedge-shaped lobules by connective tissue called septa. Coiled in each wedge are seminiferous tubules that produce sperm.
Sperm are immobile at body temperature; therefore, the scrotum and penis are external to the body, as illustrated in Figure \(1\) so that a proper temperature is maintained for motility. In land mammals, the pair of testes must be suspended outside the body at about 2° C lower than body temperature to produce viable sperm. Infertility can occur in land mammals when the testes do not descend through the abdominal cavity during fetal development.
Art Connection
Which of the following statements about the male reproductive system is false?
1. The vas deferens carries sperm from the testes to the penis.
2. Sperm mature in seminiferous tubules in the testes.
3. Both the prostate and the bulbourethral glands produce components of the semen.
4. The prostate gland is located in the testes.
Sperm mature in seminiferous tubules that are coiled inside the testes, as illustrated in Figure \(1\). The walls of the seminiferous tubules are made up of the developing sperm cells, with the least developed sperm at the periphery of the tubule and the fully developed sperm in the lumen. The sperm cells are mixed with “nursemaid” cells called Sertoli cells which protect the germ cells and promote their development. Other cells mixed in the wall of the tubules are the interstitial cells of Leydig. These cells produce high levels of testosterone once the male reaches adolescence.
When the sperm have developed flagella and are nearly mature, they leave the testicles and enter the epididymis, shown in Figure \(1\). This structure resembles a comma and lies along the top and posterior portion of the testes; it is the site of sperm maturation. The sperm leave the epididymis and enter the vas deferens (or ductus deferens), which carries the sperm, behind the bladder, and forms the ejaculatory duct with the duct from the seminal vesicles. During a vasectomy, a section of the vas deferens is removed, preventing sperm from being passed out of the body during ejaculation and preventing fertilization.
Semen is a mixture of sperm and spermatic duct secretions (about 10 percent of the total) and fluids from accessory glands that contribute most of the semen’s volume. Sperm are haploid cells, consisting of a flagellum as a tail, a neck that contains the cell’s energy-producing mitochondria, and a head that contains the genetic material. Figure \(2\) shows a micrograph of human sperm as well as a diagram of the parts of the sperm. An acrosome is found at the top of the head of the sperm. This structure contains lysosomal enzymes that can digest the protective coverings that surround the egg to help the sperm penetrate and fertilize the egg. An ejaculate will contain from two to five milliliters of fluid with from 50–120 million sperm per milliliter.
The bulk of the semen comes from the accessory glands associated with the male reproductive system. These are the seminal vesicles, the prostate gland, and the bulbourethral gland, all of which are illustrated in Figure \(1\). The seminal vesicles are a pair of glands that lie along the posterior border of the urinary bladder. The glands make a solution that is thick, yellowish, and alkaline. As sperm are only motile in an alkaline environment, a basic pH is important to reverse the acidity of the vaginal environment. The solution also contains mucus, fructose (a sperm mitochondrial nutrient), a coagulating enzyme, ascorbic acid, and local-acting hormones called prostaglandins. The seminal vesicle glands account for 60 percent of the bulk of semen.
The penis, illustrated in Figure \(1\), is an organ that drains urine from the renal bladder and functions as a copulatory organ during intercourse. The penis contains three tubes of erectile tissue running through the length of the organ. These consist of a pair of tubes on the dorsal side, called the corpus cavernosum, and a single tube of tissue on the ventral side, called the corpus spongiosum. This tissue will become engorged with blood, becoming erect and hard, in preparation for intercourse. The organ is inserted into the vagina culminating with an ejaculation. During intercourse, the smooth muscle sphincters at the opening to the renal bladder close and prevent urine from entering the penis. An orgasm is a two-stage process: first, glands and accessory organs connected to the testes contract, then semen (containing sperm) is expelled through the urethra during ejaculation. After intercourse, the blood drains from the erectile tissue and the penis becomes flaccid.
The walnut-shaped prostate gland surrounds the urethra, the connection to the urinary bladder. It has a series of short ducts that directly connect to the urethra. The gland is a mixture of smooth muscle and glandular tissue. The muscle provides much of the force needed for ejaculation to occur. The glandular tissue makes a thin, milky fluid that contains citrate (a nutrient), enzymes, and prostate specific antigen (PSA). PSA is a proteolytic enzyme that helps to liquefy the ejaculate several minutes after release from the male. Prostate gland secretions account for about 30 percent of the bulk of semen.
The bulbourethral gland, or Cowper’s gland, releases its secretion prior to the release of the bulk of the semen. It neutralizes any acid residue in the urethra left over from urine. This usually accounts for a couple of drops of fluid in the total ejaculate and may contain a few sperm. Withdrawal of the penis from the vagina before ejaculation to prevent pregnancy may not work if sperm are present in the bulbourethral gland secretions. The location and functions of the male reproductive organs are summarized in Table \(1\).
Table \(1\): Male Reproductive Anatomy
Organ Location Function
Scrotum External Carry and support testes
Penis External Deliver urine, copulating organ
Testes Internal Produce sperm and male hormones
Seminal Vesicles Internal Contribute to semen production
Prostate Gland Internal Contribute to semen production
Bulbourethral Glands Internal Clean urethra at ejaculation
Female Reproductive Anatomy
A number of reproductive structures are exterior to the female’s body. These include the breasts and the vulva, which consists of the mons pubis, clitoris, labia majora, labia minora, and the vestibular glands, all illustrated in Figure \(3\). The location and functions of the female reproductive organs are summarized in Table \(2\). The vulva is an area associated with the vestibule which includes the structures found in the inguinal (groin) area of women. The mons pubis is a round, fatty area that overlies the pubic symphysis. The clitoris is a structure with erectile tissue that contains a large number of sensory nerves and serves as a source of stimulation during intercourse. The labia majora are a pair of elongated folds of tissue that run posterior from the mons pubis and enclose the other components of the vulva. The labia majora derive from the same tissue that produces the scrotum in a male. The labia minora are thin folds of tissue centrally located within the labia majora. These labia protect the openings to the vagina and urethra. The mons pubis and the anterior portion of the labia majora become covered with hair during adolescence; the labia minora is hairless. The greater vestibular glands are found at the sides of the vaginal opening and provide lubrication during intercourse.
Table \(2\): Female Reproductive Anatomy
Organ Location Function
Clitoris External Sensory organ
Mons pubis External Fatty area overlying pubic bone
Labia majora External Covers labia minora
Labia minora External Covers vestibule
Greater vestibular glands External Secrete mucus; lubricate vagina
Breast External Produce and deliver milk
Ovaries Internal Carry and develop eggs
Oviducts (Fallopian tubes) Internal Transport egg to uterus
Uterus Internal Support developing embryo
Vagina Internal Common tube for intercourse, birth canal, passing menstrual flow
The breasts consist of mammary glands and fat. The size of the breast is determined by the amount of fat deposited behind the gland. Each gland consists of 15 to 25 lobes that have ducts that empty at the nipple and that supply the nursing child with nutrient- and antibody-rich milk to aid development and protect the child.
Internal female reproductive structures include ovaries, oviducts, the uterus, and the vagina, shown in Figure \(3\). The pair of ovaries is held in place in the abdominal cavity by a system of ligaments. Ovaries consist of a medulla and cortex: the medulla contains nerves and blood vessels to supply the cortex with nutrients and remove waste. The outer layers of cells of the cortex are the functional parts of the ovaries. The cortex is made up of follicular cells that surround eggs that develop during fetal development in utero. During the menstrual period, a batch of follicular cells develops and prepares the eggs for release. At ovulation, one follicle ruptures and one egg is released, as illustrated in Figure \(4\).
The oviducts, or fallopian tubes, extend from the uterus in the lower abdominal cavity to the ovaries, but they are not in contact with the ovaries. The lateral ends of the oviducts flare out into a trumpet-like structure and have a fringe of finger-like projections called fimbriae, illustrated in Figure \(4\). When an egg is released at ovulation, the fimbrae help the non-motile egg enter into the tube and passage to the uterus. The walls of the oviducts are ciliated and are made up mostly of smooth muscle. The cilia beat toward the middle, and the smooth muscle contracts in the same direction, moving the egg toward the uterus. Fertilization usually takes place within the oviducts and the developing embryo is moved toward the uterus for development. It usually takes the egg or embryo a week to travel through the oviduct. Sterilization in women is called a tubal ligation; it is analogous to a vasectomy in males in that the oviducts are severed and sealed.
The uterus is a structure about the size of a woman’s fist. This is lined with an endometrium rich in blood vessels and mucus glands. The uterus supports the developing embryo and fetus during gestation. The thickest portion of the wall of the uterus is made of smooth muscle. Contractions of the smooth muscle in the uterus aid in passing the baby through the vagina during labor. A portion of the lining of the uterus sloughs off during each menstrual period, and then builds up again in preparation for an implantation. Part of the uterus, called the cervix, protrudes into the top of the vagina. The cervix functions as the birth canal.
The vagina is a muscular tube that serves several purposes. It allows menstrual flow to leave the body. It is the receptacle for the penis during intercourse and the vessel for the delivery of offspring. It is lined by stratified squamous epithelial cells to protect the underlying tissue.
Sexual Response during Intercourse
The sexual response in humans is both psychological and physiological. Both sexes experience sexual arousal through psychological and physical stimulation. There are four phases of the sexual response. During phase one, called excitement, vasodilation leads to vasocongestion in erectile tissues in both men and women. The nipples, clitoris, labia, and penis engorge with blood and become enlarged. Vaginal secretions are released to lubricate the vagina to facilitate intercourse. During the second phase, called the plateau, stimulation continues, the outer third of the vaginal wall enlarges with blood, and breathing and heart rate increase.
During phase three, or orgasm, rhythmic, involuntary contractions of muscles occur in both sexes. In the male, the reproductive accessory glands and tubules constrict placing semen in the urethra, then the urethra contracts expelling the semen through the penis. In women, the uterus and vaginal muscles contract in waves that may last slightly less than a second each. During phase four, or resolution, the processes described in the first three phases reverse themselves and return to their normal state. Men experience a refractory period in which they cannot maintain an erection or ejaculate for a period of time ranging from minutes to hours.
Gametogenesis (Spermatogenesis and Oogenesis)
Gametogenesis, the production of sperm and eggs, takes place through the process of meiosis. During meiosis, two cell divisions separate the paired chromosomes in the nucleus and then separate the chromatids that were made during an earlier stage of the cell’s life cycle. Meiosis produces haploid cells with half of each pair of chromosomes normally found in diploid cells. The production of sperm is called spermatogenesis and the production of eggs is called oogenesis.
Spermatogenesis
Spermatogenesis, illustrated in Figure \(5\), occurs in the wall of the seminiferous tubules, with stem cells at the periphery of the tube and the spermatozoa at the lumen of the tube. Immediately under the capsule of the tubule are diploid, undifferentiated cells. These stem cells, called spermatogonia (singular: spermatagonium), go through mitosis with one offspring going on to differentiate into a sperm cell and the other giving rise to the next generation of sperm.
Meiosis starts with a cell called a primary spermatocyte. At the end of the first meiotic division, a haploid cell is produced called a secondary spermatocyte. This cell is haploid and must go through another meiotic cell division. The cell produced at the end of meiosis is called a spermatid and when it reaches the lumen of the tubule and grows a flagellum, it is called a sperm cell. Four sperm result from each primary spermatocyte that goes through meiosis.
Stem cells are deposited during gestation and are present at birth through the beginning of adolescence, but in an inactive state. During adolescence, gonadotropic hormones from the anterior pituitary cause the activation of these cells and the production of viable sperm. This continues into old age.
Link to Learning
Visit this site to see the process of spermatogenesis.
Oogenesis
Oogenesis, illustrated in Figure \(6\), occurs in the outermost layers of the ovaries. As with sperm production, oogenesis starts with a germ cell, called an oogonium (plural: oogonia), but this cell undergoes mitosis to increase in number, eventually resulting in up to about one to two million cells in the embryo.
The cell starting meiosis is called a primary oocyte, as shown in Figure \(6\). This cell will start the first meiotic division and be arrested in its progress in the first prophase stage. At the time of birth, all future eggs are in the prophase stage. At adolescence, anterior pituitary hormones cause the development of a number of follicles in an ovary. This results in the primary oocyte finishing the first meiotic division. The cell divides unequally, with most of the cellular material and organelles going to one cell, called a secondary oocyte, and only one set of chromosomes and a small amount of cytoplasm going to the other cell. This second cell is called a polar body and usually dies. A secondary meiotic arrest occurs, this time at the metaphase II stage. At ovulation, this secondary oocyte will be released and travel toward the uterus through the oviduct. If the secondary oocyte is fertilized, the cell continues through the meiosis II, producing a second polar body and a fertilized egg containing all 46 chromosomes of a human being, half of them coming from the sperm.
Egg production begins before birth, is arrested during meiosis until puberty, and then individual cells continue through at each menstrual cycle. One egg is produced from each meiotic process, with the extra chromosomes and chromatids going into polar bodies that degenerate and are reabsorbed by the body.
Summary
As animals became more complex, specific organs and organ systems developed to support specific functions for the organism. The reproductive structures that evolved in land animals allow males and females to mate, fertilize internally, and support the growth and development of offspring. Processes developed to produce reproductive cells that had exactly half the number of chromosomes of each parent so that new combinations would have the appropriate amount of genetic material. Gametogenesis, the production of sperm (spermatogenesis) and eggs (oogenesis), takes place through the process of meiosis.
Figure \(1\): Which of the following statements about the male reproductive system is false?
1. The vas deferens carries sperm from the testes to the penis.
2. Sperm mature in seminiferous tubules in the testes.
3. Both the prostate and the bulbourethral glands produce components of the semen.
4. The prostate gland is located in the testes.
Answer
D
Glossary
bulbourethral gland
secretion that cleanses the urethra prior to ejaculation
clitoris
sensory structure in females; stimulated during sexual arousal
labia majora
large folds of tissue covering the inguinal area
labia minora
smaller folds of tissue within the labia majora
oogenesis
process of producing haploid eggs
oviduct
(also, fallopian tube) muscular tube connecting the uterus with the ovary area
penis
male reproductive structure for urine elimination and copulation
prostate gland
structure that is a mixture of smooth muscle and glandular material and that contributes to semen
scrotum
sac containing testes; exterior to the body
semen
fluid mixture of sperm and supporting materials
seminal vesicle
secretory accessory gland in males; contributes to semen
seminiferous tubule
site of sperm production in testes
spermatogenesis
process of producing haploid sperm
testes
pair of reproductive organs in males
uterus
environment for developing embryo and fetus
vagina
muscular tube for the passage of menstrual flow, copulation, and birth of offspring
51.03: Structure and Function of the Human Male Reproductive System
Skills to Develop
• Describe the roles of male and female reproductive hormones
• Discuss the interplay of the ovarian and menstrual cycles
• Describe the process of menopause
The human male and female reproductive cycles are controlled by the interaction of hormones from the hypothalamus and anterior pituitary with hormones from reproductive tissues and organs. In both sexes, the hypothalamus monitors and causes the release of hormones from the pituitary gland. When the reproductive hormone is required, the hypothalamus sends a gonadotropin-releasing hormone (GnRH) to the anterior pituitary. This causes the release of follicle stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary into the blood. Note that the body must reach puberty for the adrenals to release the hormones that must be present for GnRH to be produced. Although FSH and LH are named after their functions in female reproduction, they are produced in both sexes and play important roles in controlling reproduction. Other hormones have specific functions in the male and female reproductive systems.
Male Hormones
At the onset of puberty, the hypothalamus causes the release of FSH and LH into the male system for the first time. FSH enters the testes and stimulates the Sertoli cells to begin facilitating spermatogenesis using negative feedback, as illustrated in Figure \(1\). LH also enters the testes and stimulates the interstitial cells of Leydig to make and release testosterone into the testes and the blood.
Testosterone, the hormone responsible for the secondary sexual characteristics that develop in the male during adolescence, stimulates spermatogenesis. These secondary sex characteristics include a deepening of the voice, the growth of facial, axillary, and pubic hair, and the beginnings of the sex drive.
A negative feedback system occurs in the male with rising levels of testosterone acting on the hypothalamus and anterior pituitary to inhibit the release of GnRH, FSH, and LH. The Sertoli cells produce the hormone inhibin, which is released into the blood when the sperm count is too high. This inhibits the release of GnRH and FSH, which will cause spermatogenesis to slow down. If the sperm count reaches 20 million/ml, the Sertoli cells cease the release of inhibin, and the sperm count increases.
Female Hormones
The control of reproduction in females is more complex. As with the male, the anterior pituitary hormones cause the release of the hormones FSH and LH. In addition, estrogens and progesterone are released from the developing follicles. Estrogen is the reproductive hormone in females that assists in endometrial regrowth, ovulation, and calcium absorption; it is also responsible for the secondary sexual characteristics of females. These include breast development, flaring of the hips, and a shorter period necessary for bone maturation. Progesterone assists in endometrial re-growth and inhibition of FSH and LH release.
In females, FSH stimulates development of egg cells, called ova, which develop in structures called follicles. Follicle cells produce the hormone inhibin, which inhibits FSH production. LH also plays a role in the development of ova, induction of ovulation, and stimulation of estradiol and progesterone production by the ovaries. Estradiol and progesterone are steroid hormones that prepare the body for pregnancy. Estradiol produces secondary sex characteristics in females, while both estradiol and progesterone regulate the menstrual cycle.
The Ovarian Cycle and the Menstrual Cycle
The ovarian cycle governs the preparation of endocrine tissues and release of eggs, while the menstrual cycle governs the preparation and maintenance of the uterine lining. These cycles occur concurrently and are coordinated over a 22–32 day cycle, with an average length of 28 days.
The first half of the ovarian cycle is the follicular phase shown in Figure \(2\). Slowly rising levels of FSH and LH cause the growth of follicles on the surface of the ovary. This process prepares the egg for ovulation. As the follicles grow, they begin releasing estrogens and a low level of progesterone. Progesterone maintains the endometrium to help ensure pregnancy. The trip through the fallopian tube takes about seven days. At this stage of development, called the morula, there are 30-60 cells. If pregnancy implantation does not occur, the lining is sloughed off. After about five days, estrogen levels rise and the menstrual cycle enters the proliferative phase. The endometrium begins to regrow, replacing the blood vessels and glands that deteriorated during the end of the last cycle.
Art Connection
Which of the following statements about hormone regulation of the female reproductive cycle is false?
1. LH and FSH are produced in the pituitary, and estradiol and progesterone are produced in the ovaries.
2. Estradiol and progesterone secreted from the corpus luteum cause the endometrium to thicken.
3. Both progesterone and estradiol are produced by the follicles.
4. Secretion of GnRH by the hypothalamus is inhibited by low levels of estradiol but stimulated by high levels of estradiol.
Just prior to the middle of the cycle (approximately day 14), the high level of estrogen causes FSH and especially LH to rise rapidly, then fall. The spike in LH causes ovulation: the most mature follicle, like that shown in Figure \(3\), ruptures and releases its egg. The follicles that did not rupture degenerate and their eggs are lost. The level of estrogen decreases when the extra follicles degenerate.
Following ovulation, the ovarian cycle enters its luteal phase, illustrated in Figure \(4\) and the menstrual cycle enters its secretory phase, both of which run from about day 15 to 28. The luteal and secretory phases refer to changes in the ruptured follicle. The cells in the follicle undergo physical changes and produce a structure called a corpus luteum. The corpus luteum produces estrogen and progesterone. The progesterone facilitates the regrowth of the uterine lining and inhibits the release of further FSH and LH. The uterus is being prepared to accept a fertilized egg, should it occur during this cycle. The inhibition of FSH and LH prevents any further eggs and follicles from developing, while the progesterone is elevated. The level of estrogen produced by the corpus luteum increases to a steady level for the next few days.
If no fertilized egg is implanted into the uterus, the corpus luteum degenerates and the levels of estrogen and progesterone decrease. The endometrium begins to degenerate as the progesterone levels drop, initiating the next menstrual cycle. The decrease in progesterone also allows the hypothalamus to send GnRH to the anterior pituitary, releasing FSH and LH and starting the cycles again. Figure \(4\) visually compares the ovarian and uterine cycles as well as the commensurate hormone levels.
Art Connection
Which of the following statements about the menstrual cycle is false?
1. Progesterone levels rise during the luteal phase of the ovarian cycle and the secretory phase of the uterine cycle.
2. Menstruation occurs just after LH and FSH levels peak.
3. Menstruation occurs after progesterone levels drop.
4. Estrogen levels rise before ovulation, while progesterone levels rise after.
Menopause
As women approach their mid-40s to mid-50s, their ovaries begin to lose their sensitivity to FSH and LH. Menstrual periods become less frequent and finally cease; this is menopause. There are still eggs and potential follicles on the ovaries, but without the stimulation of FSH and LH, they will not produce a viable egg to be released. The outcome of this is the inability to have children.
The side effects of menopause include hot flashes, heavy sweating (especially at night), headaches, some hair loss, muscle pain, vaginal dryness, insomnia, depression, weight gain, and mood swings. Estrogen is involved in calcium metabolism and, without it, blood levels of calcium decrease. To replenish the blood, calcium is lost from bone which may decrease the bone density and lead to osteoporosis. Supplementation of estrogen in the form of hormone replacement therapy (HRT) can prevent bone loss, but the therapy can have negative side effects. While HRT is thought to give some protection from colon cancer, osteoporosis, heart disease, macular degeneration, and possibly depression, its negative side effects include increased risk of: stroke or heart attack, blood clots, breast cancer, ovarian cancer, endometrial cancer, gall bladder disease, and possibly dementia.
Career Connection: Reproductive Endocrinologist
A reproductive endocrinologist is a physician who treats a variety of hormonal disorders related to reproduction and infertility in both men and women. The disorders include menstrual problems, infertility, pregnancy loss, sexual dysfunction, and menopause. Doctors may use fertility drugs, surgery, or assisted reproductive techniques (ART) in their therapy. ART involves the use of procedures to manipulate the egg or sperm to facilitate reproduction, such as in vitro fertilization.
Reproductive endocrinologists undergo extensive medical training, first in a four-year residency in obstetrics and gynecology, then in a three-year fellowship in reproductive endocrinology. To be board certified in this area, the physician must pass written and oral exams in both areas.
Summary
The male and female reproductive cycles are controlled by hormones released from the hypothalamus and anterior pituitary as well as hormones from reproductive tissues and organs. The hypothalamus monitors the need for the FSH and LH hormones made and released from the anterior pituitary. FSH and LH affect reproductive structures to cause the formation of sperm and the preparation of eggs for release and possible fertilization. In the male, FSH and LH stimulate Sertoli cells and interstitial cells of Leydig in the testes to facilitate sperm production. The Leydig cells produce testosterone, which also is responsible for the secondary sexual characteristics of males. In females, FSH and LH cause estrogen and progesterone to be produced. They regulate the female reproductive system which is divided into the ovarian cycle and the menstrual cycle. Menopause occurs when the ovaries lose their sensitivity to FSH and LH and the female reproductive cycles slow to a stop.
Art Connections
Figure \(2\): Which of the following statements about hormone regulation of the female reproductive cycle is false?
1. LH and FSH are produced in the pituitary, and estradiol and progesterone are produced in the ovaries.
2. Estradiol and progesterone secreted from the corpus luteum cause the endometrium to thicken.
3. Both progesterone and estradiol are produced by the follicles.
4. Secretion of GnRH by the hypothalamus is inhibited by low levels of estradiol but stimulated by high levels of estradiol.
Answer
C
Figure \(4\): Which of the following statements about the menstrual cycle is false?
1. Progesterone levels rise during the luteal phase of the ovarian cycle and the secretory phase of the uterine cycle.
2. Menstruation occurs just after LH and FSH levels peak.
3. Menstruation occurs after progesterone levels drop.
4. Estrogen levels rise before ovulation, while progesterone levels rise after.
Answer
B
Glossary
estrogen
reproductive hormone in females that assists in endometrial regrowth, ovulation, and calcium absorption
follicle stimulating hormone (FSH)
reproductive hormone that causes sperm production in men and follicle development in women
gonadotropin-releasing hormone (GnRH)
hormone from the hypothalamus that causes the release of FSH and LH from the anterior pituitary
inhibin
hormone made by Sertoli cells; provides negative feedback to hypothalamus in control of FSH and GnRH release
interstitial cell of Leydig
cell in seminiferous tubules that makes testosterone
luteinizing hormone (LH)
reproductive hormone in both men and women, causes testosterone production in men and ovulation and lactation in women
menopause
loss of reproductive capacity in women due to decreased sensitivity of the ovaries to FSH and LH
menstrual cycle
cycle of the degradation and re-growth of the endometrium
ovarian cycle
cycle of preparation of egg for ovulation and the conversion of the follicle to the corpus luteum
ovulation
release of the egg by the most mature follicle
progesterone
reproductive hormone in women; assists in endometrial re-growth and inhibition of FSH and LH release
Sertoli cell
cell in seminiferous tubules that assists developing sperm and makes inhibin
testosterone
reproductive hormone in men that assists in sperm production and promoting secondary sexual characteristics | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/51%3A_The_Reproductive_System/51.03%3A_Structure_and_Function_of_the_Human_Male_Reproductive_System/51.3.01%3A_Hormonal_Control_of_Human_Reproduction.txt |
Skills to Develop
• Describe human male and female reproductive anatomies
• Discuss the human sexual response
• Describe spermatogenesis and oogenesis and discuss their differences and similarities
As animals became more complex, specific organs and organ systems developed to support specific functions for the organism. The reproductive structures that evolved in land animals allow males and females to mate, fertilize internally, and support the growth and development of offspring.
Human Reproductive Anatomy
The reproductive tissues of male and female humans develop similarly in utero until a low level of the hormone testosterone is released from male gonads. Testosterone causes the undeveloped tissues to differentiate into male sexual organs. When testosterone is absent, the tissues develop into female sexual tissues. Primitive gonads become testes or ovaries. Tissues that produce a penis in males produce a clitoris in females. The tissue that will become the scrotum in a male becomes the labia in a female; that is, they are homologous structures.
Male Reproductive Anatomy
In the male reproductive system, the scrotum houses the testicles or testes (singular: testis), including providing passage for blood vessels, nerves, and muscles related to testicular function. The testes are a pair of male reproductive organs that produce sperm and some reproductive hormones. Each testis is approximately 2.5 by 3.8 cm (1.5 by 1 in) in size and divided into wedge-shaped lobules by connective tissue called septa. Coiled in each wedge are seminiferous tubules that produce sperm.
Sperm are immobile at body temperature; therefore, the scrotum and penis are external to the body, as illustrated in Figure \(1\) so that a proper temperature is maintained for motility. In land mammals, the pair of testes must be suspended outside the body at about 2° C lower than body temperature to produce viable sperm. Infertility can occur in land mammals when the testes do not descend through the abdominal cavity during fetal development.
Art Connection
Which of the following statements about the male reproductive system is false?
1. The vas deferens carries sperm from the testes to the penis.
2. Sperm mature in seminiferous tubules in the testes.
3. Both the prostate and the bulbourethral glands produce components of the semen.
4. The prostate gland is located in the testes.
Sperm mature in seminiferous tubules that are coiled inside the testes, as illustrated in Figure \(1\). The walls of the seminiferous tubules are made up of the developing sperm cells, with the least developed sperm at the periphery of the tubule and the fully developed sperm in the lumen. The sperm cells are mixed with “nursemaid” cells called Sertoli cells which protect the germ cells and promote their development. Other cells mixed in the wall of the tubules are the interstitial cells of Leydig. These cells produce high levels of testosterone once the male reaches adolescence.
When the sperm have developed flagella and are nearly mature, they leave the testicles and enter the epididymis, shown in Figure \(1\). This structure resembles a comma and lies along the top and posterior portion of the testes; it is the site of sperm maturation. The sperm leave the epididymis and enter the vas deferens (or ductus deferens), which carries the sperm, behind the bladder, and forms the ejaculatory duct with the duct from the seminal vesicles. During a vasectomy, a section of the vas deferens is removed, preventing sperm from being passed out of the body during ejaculation and preventing fertilization.
Semen is a mixture of sperm and spermatic duct secretions (about 10 percent of the total) and fluids from accessory glands that contribute most of the semen’s volume. Sperm are haploid cells, consisting of a flagellum as a tail, a neck that contains the cell’s energy-producing mitochondria, and a head that contains the genetic material. Figure \(2\) shows a micrograph of human sperm as well as a diagram of the parts of the sperm. An acrosome is found at the top of the head of the sperm. This structure contains lysosomal enzymes that can digest the protective coverings that surround the egg to help the sperm penetrate and fertilize the egg. An ejaculate will contain from two to five milliliters of fluid with from 50–120 million sperm per milliliter.
The bulk of the semen comes from the accessory glands associated with the male reproductive system. These are the seminal vesicles, the prostate gland, and the bulbourethral gland, all of which are illustrated in Figure \(1\). The seminal vesicles are a pair of glands that lie along the posterior border of the urinary bladder. The glands make a solution that is thick, yellowish, and alkaline. As sperm are only motile in an alkaline environment, a basic pH is important to reverse the acidity of the vaginal environment. The solution also contains mucus, fructose (a sperm mitochondrial nutrient), a coagulating enzyme, ascorbic acid, and local-acting hormones called prostaglandins. The seminal vesicle glands account for 60 percent of the bulk of semen.
The penis, illustrated in Figure \(1\), is an organ that drains urine from the renal bladder and functions as a copulatory organ during intercourse. The penis contains three tubes of erectile tissue running through the length of the organ. These consist of a pair of tubes on the dorsal side, called the corpus cavernosum, and a single tube of tissue on the ventral side, called the corpus spongiosum. This tissue will become engorged with blood, becoming erect and hard, in preparation for intercourse. The organ is inserted into the vagina culminating with an ejaculation. During intercourse, the smooth muscle sphincters at the opening to the renal bladder close and prevent urine from entering the penis. An orgasm is a two-stage process: first, glands and accessory organs connected to the testes contract, then semen (containing sperm) is expelled through the urethra during ejaculation. After intercourse, the blood drains from the erectile tissue and the penis becomes flaccid.
The walnut-shaped prostate gland surrounds the urethra, the connection to the urinary bladder. It has a series of short ducts that directly connect to the urethra. The gland is a mixture of smooth muscle and glandular tissue. The muscle provides much of the force needed for ejaculation to occur. The glandular tissue makes a thin, milky fluid that contains citrate (a nutrient), enzymes, and prostate specific antigen (PSA). PSA is a proteolytic enzyme that helps to liquefy the ejaculate several minutes after release from the male. Prostate gland secretions account for about 30 percent of the bulk of semen.
The bulbourethral gland, or Cowper’s gland, releases its secretion prior to the release of the bulk of the semen. It neutralizes any acid residue in the urethra left over from urine. This usually accounts for a couple of drops of fluid in the total ejaculate and may contain a few sperm. Withdrawal of the penis from the vagina before ejaculation to prevent pregnancy may not work if sperm are present in the bulbourethral gland secretions. The location and functions of the male reproductive organs are summarized in Table \(1\).
Table \(1\): Male Reproductive Anatomy
Organ Location Function
Scrotum External Carry and support testes
Penis External Deliver urine, copulating organ
Testes Internal Produce sperm and male hormones
Seminal Vesicles Internal Contribute to semen production
Prostate Gland Internal Contribute to semen production
Bulbourethral Glands Internal Clean urethra at ejaculation
Female Reproductive Anatomy
A number of reproductive structures are exterior to the female’s body. These include the breasts and the vulva, which consists of the mons pubis, clitoris, labia majora, labia minora, and the vestibular glands, all illustrated in Figure \(3\). The location and functions of the female reproductive organs are summarized in Table \(2\). The vulva is an area associated with the vestibule which includes the structures found in the inguinal (groin) area of women. The mons pubis is a round, fatty area that overlies the pubic symphysis. The clitoris is a structure with erectile tissue that contains a large number of sensory nerves and serves as a source of stimulation during intercourse. The labia majora are a pair of elongated folds of tissue that run posterior from the mons pubis and enclose the other components of the vulva. The labia majora derive from the same tissue that produces the scrotum in a male. The labia minora are thin folds of tissue centrally located within the labia majora. These labia protect the openings to the vagina and urethra. The mons pubis and the anterior portion of the labia majora become covered with hair during adolescence; the labia minora is hairless. The greater vestibular glands are found at the sides of the vaginal opening and provide lubrication during intercourse.
Table \(2\): Female Reproductive Anatomy
Organ Location Function
Clitoris External Sensory organ
Mons pubis External Fatty area overlying pubic bone
Labia majora External Covers labia minora
Labia minora External Covers vestibule
Greater vestibular glands External Secrete mucus; lubricate vagina
Breast External Produce and deliver milk
Ovaries Internal Carry and develop eggs
Oviducts (Fallopian tubes) Internal Transport egg to uterus
Uterus Internal Support developing embryo
Vagina Internal Common tube for intercourse, birth canal, passing menstrual flow
The breasts consist of mammary glands and fat. The size of the breast is determined by the amount of fat deposited behind the gland. Each gland consists of 15 to 25 lobes that have ducts that empty at the nipple and that supply the nursing child with nutrient- and antibody-rich milk to aid development and protect the child.
Internal female reproductive structures include ovaries, oviducts, the uterus, and the vagina, shown in Figure \(3\). The pair of ovaries is held in place in the abdominal cavity by a system of ligaments. Ovaries consist of a medulla and cortex: the medulla contains nerves and blood vessels to supply the cortex with nutrients and remove waste. The outer layers of cells of the cortex are the functional parts of the ovaries. The cortex is made up of follicular cells that surround eggs that develop during fetal development in utero. During the menstrual period, a batch of follicular cells develops and prepares the eggs for release. At ovulation, one follicle ruptures and one egg is released, as illustrated in Figure \(4\).
The oviducts, or fallopian tubes, extend from the uterus in the lower abdominal cavity to the ovaries, but they are not in contact with the ovaries. The lateral ends of the oviducts flare out into a trumpet-like structure and have a fringe of finger-like projections called fimbriae, illustrated in Figure \(4\). When an egg is released at ovulation, the fimbrae help the non-motile egg enter into the tube and passage to the uterus. The walls of the oviducts are ciliated and are made up mostly of smooth muscle. The cilia beat toward the middle, and the smooth muscle contracts in the same direction, moving the egg toward the uterus. Fertilization usually takes place within the oviducts and the developing embryo is moved toward the uterus for development. It usually takes the egg or embryo a week to travel through the oviduct. Sterilization in women is called a tubal ligation; it is analogous to a vasectomy in males in that the oviducts are severed and sealed.
The uterus is a structure about the size of a woman’s fist. This is lined with an endometrium rich in blood vessels and mucus glands. The uterus supports the developing embryo and fetus during gestation. The thickest portion of the wall of the uterus is made of smooth muscle. Contractions of the smooth muscle in the uterus aid in passing the baby through the vagina during labor. A portion of the lining of the uterus sloughs off during each menstrual period, and then builds up again in preparation for an implantation. Part of the uterus, called the cervix, protrudes into the top of the vagina. The cervix functions as the birth canal.
The vagina is a muscular tube that serves several purposes. It allows menstrual flow to leave the body. It is the receptacle for the penis during intercourse and the vessel for the delivery of offspring. It is lined by stratified squamous epithelial cells to protect the underlying tissue.
Sexual Response during Intercourse
The sexual response in humans is both psychological and physiological. Both sexes experience sexual arousal through psychological and physical stimulation. There are four phases of the sexual response. During phase one, called excitement, vasodilation leads to vasocongestion in erectile tissues in both men and women. The nipples, clitoris, labia, and penis engorge with blood and become enlarged. Vaginal secretions are released to lubricate the vagina to facilitate intercourse. During the second phase, called the plateau, stimulation continues, the outer third of the vaginal wall enlarges with blood, and breathing and heart rate increase.
During phase three, or orgasm, rhythmic, involuntary contractions of muscles occur in both sexes. In the male, the reproductive accessory glands and tubules constrict placing semen in the urethra, then the urethra contracts expelling the semen through the penis. In women, the uterus and vaginal muscles contract in waves that may last slightly less than a second each. During phase four, or resolution, the processes described in the first three phases reverse themselves and return to their normal state. Men experience a refractory period in which they cannot maintain an erection or ejaculate for a period of time ranging from minutes to hours.
Gametogenesis (Spermatogenesis and Oogenesis)
Gametogenesis, the production of sperm and eggs, takes place through the process of meiosis. During meiosis, two cell divisions separate the paired chromosomes in the nucleus and then separate the chromatids that were made during an earlier stage of the cell’s life cycle. Meiosis produces haploid cells with half of each pair of chromosomes normally found in diploid cells. The production of sperm is called spermatogenesis and the production of eggs is called oogenesis.
Spermatogenesis
Spermatogenesis, illustrated in Figure \(5\), occurs in the wall of the seminiferous tubules, with stem cells at the periphery of the tube and the spermatozoa at the lumen of the tube. Immediately under the capsule of the tubule are diploid, undifferentiated cells. These stem cells, called spermatogonia (singular: spermatagonium), go through mitosis with one offspring going on to differentiate into a sperm cell and the other giving rise to the next generation of sperm.
Meiosis starts with a cell called a primary spermatocyte. At the end of the first meiotic division, a haploid cell is produced called a secondary spermatocyte. This cell is haploid and must go through another meiotic cell division. The cell produced at the end of meiosis is called a spermatid and when it reaches the lumen of the tubule and grows a flagellum, it is called a sperm cell. Four sperm result from each primary spermatocyte that goes through meiosis.
Stem cells are deposited during gestation and are present at birth through the beginning of adolescence, but in an inactive state. During adolescence, gonadotropic hormones from the anterior pituitary cause the activation of these cells and the production of viable sperm. This continues into old age.
Link to Learning
Visit this site to see the process of spermatogenesis.
Oogenesis
Oogenesis, illustrated in Figure \(6\), occurs in the outermost layers of the ovaries. As with sperm production, oogenesis starts with a germ cell, called an oogonium (plural: oogonia), but this cell undergoes mitosis to increase in number, eventually resulting in up to about one to two million cells in the embryo.
The cell starting meiosis is called a primary oocyte, as shown in Figure \(6\). This cell will start the first meiotic division and be arrested in its progress in the first prophase stage. At the time of birth, all future eggs are in the prophase stage. At adolescence, anterior pituitary hormones cause the development of a number of follicles in an ovary. This results in the primary oocyte finishing the first meiotic division. The cell divides unequally, with most of the cellular material and organelles going to one cell, called a secondary oocyte, and only one set of chromosomes and a small amount of cytoplasm going to the other cell. This second cell is called a polar body and usually dies. A secondary meiotic arrest occurs, this time at the metaphase II stage. At ovulation, this secondary oocyte will be released and travel toward the uterus through the oviduct. If the secondary oocyte is fertilized, the cell continues through the meiosis II, producing a second polar body and a fertilized egg containing all 46 chromosomes of a human being, half of them coming from the sperm.
Egg production begins before birth, is arrested during meiosis until puberty, and then individual cells continue through at each menstrual cycle. One egg is produced from each meiotic process, with the extra chromosomes and chromatids going into polar bodies that degenerate and are reabsorbed by the body.
Summary
As animals became more complex, specific organs and organ systems developed to support specific functions for the organism. The reproductive structures that evolved in land animals allow males and females to mate, fertilize internally, and support the growth and development of offspring. Processes developed to produce reproductive cells that had exactly half the number of chromosomes of each parent so that new combinations would have the appropriate amount of genetic material. Gametogenesis, the production of sperm (spermatogenesis) and eggs (oogenesis), takes place through the process of meiosis.
Figure \(1\): Which of the following statements about the male reproductive system is false?
1. The vas deferens carries sperm from the testes to the penis.
2. Sperm mature in seminiferous tubules in the testes.
3. Both the prostate and the bulbourethral glands produce components of the semen.
4. The prostate gland is located in the testes.
Answer
D
Glossary
bulbourethral gland
secretion that cleanses the urethra prior to ejaculation
clitoris
sensory structure in females; stimulated during sexual arousal
labia majora
large folds of tissue covering the inguinal area
labia minora
smaller folds of tissue within the labia majora
oogenesis
process of producing haploid eggs
oviduct
(also, fallopian tube) muscular tube connecting the uterus with the ovary area
penis
male reproductive structure for urine elimination and copulation
prostate gland
structure that is a mixture of smooth muscle and glandular material and that contributes to semen
scrotum
sac containing testes; exterior to the body
semen
fluid mixture of sperm and supporting materials
seminal vesicle
secretory accessory gland in males; contributes to semen
seminiferous tubule
site of sperm production in testes
spermatogenesis
process of producing haploid sperm
testes
pair of reproductive organs in males
uterus
environment for developing embryo and fetus
vagina
muscular tube for the passage of menstrual flow, copulation, and birth of offspring
51.04: Structure and Function of the Female Reproductive System
Skills to Develop
• Describe the roles of male and female reproductive hormones
• Discuss the interplay of the ovarian and menstrual cycles
• Describe the process of menopause
The human male and female reproductive cycles are controlled by the interaction of hormones from the hypothalamus and anterior pituitary with hormones from reproductive tissues and organs. In both sexes, the hypothalamus monitors and causes the release of hormones from the pituitary gland. When the reproductive hormone is required, the hypothalamus sends a gonadotropin-releasing hormone (GnRH) to the anterior pituitary. This causes the release of follicle stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary into the blood. Note that the body must reach puberty for the adrenals to release the hormones that must be present for GnRH to be produced. Although FSH and LH are named after their functions in female reproduction, they are produced in both sexes and play important roles in controlling reproduction. Other hormones have specific functions in the male and female reproductive systems.
Male Hormones
At the onset of puberty, the hypothalamus causes the release of FSH and LH into the male system for the first time. FSH enters the testes and stimulates the Sertoli cells to begin facilitating spermatogenesis using negative feedback, as illustrated in Figure \(1\). LH also enters the testes and stimulates the interstitial cells of Leydig to make and release testosterone into the testes and the blood.
Testosterone, the hormone responsible for the secondary sexual characteristics that develop in the male during adolescence, stimulates spermatogenesis. These secondary sex characteristics include a deepening of the voice, the growth of facial, axillary, and pubic hair, and the beginnings of the sex drive.
A negative feedback system occurs in the male with rising levels of testosterone acting on the hypothalamus and anterior pituitary to inhibit the release of GnRH, FSH, and LH. The Sertoli cells produce the hormone inhibin, which is released into the blood when the sperm count is too high. This inhibits the release of GnRH and FSH, which will cause spermatogenesis to slow down. If the sperm count reaches 20 million/ml, the Sertoli cells cease the release of inhibin, and the sperm count increases.
Female Hormones
The control of reproduction in females is more complex. As with the male, the anterior pituitary hormones cause the release of the hormones FSH and LH. In addition, estrogens and progesterone are released from the developing follicles. Estrogen is the reproductive hormone in females that assists in endometrial regrowth, ovulation, and calcium absorption; it is also responsible for the secondary sexual characteristics of females. These include breast development, flaring of the hips, and a shorter period necessary for bone maturation. Progesterone assists in endometrial re-growth and inhibition of FSH and LH release.
In females, FSH stimulates development of egg cells, called ova, which develop in structures called follicles. Follicle cells produce the hormone inhibin, which inhibits FSH production. LH also plays a role in the development of ova, induction of ovulation, and stimulation of estradiol and progesterone production by the ovaries. Estradiol and progesterone are steroid hormones that prepare the body for pregnancy. Estradiol produces secondary sex characteristics in females, while both estradiol and progesterone regulate the menstrual cycle.
The Ovarian Cycle and the Menstrual Cycle
The ovarian cycle governs the preparation of endocrine tissues and release of eggs, while the menstrual cycle governs the preparation and maintenance of the uterine lining. These cycles occur concurrently and are coordinated over a 22–32 day cycle, with an average length of 28 days.
The first half of the ovarian cycle is the follicular phase shown in Figure \(2\). Slowly rising levels of FSH and LH cause the growth of follicles on the surface of the ovary. This process prepares the egg for ovulation. As the follicles grow, they begin releasing estrogens and a low level of progesterone. Progesterone maintains the endometrium to help ensure pregnancy. The trip through the fallopian tube takes about seven days. At this stage of development, called the morula, there are 30-60 cells. If pregnancy implantation does not occur, the lining is sloughed off. After about five days, estrogen levels rise and the menstrual cycle enters the proliferative phase. The endometrium begins to regrow, replacing the blood vessels and glands that deteriorated during the end of the last cycle.
Art Connection
Which of the following statements about hormone regulation of the female reproductive cycle is false?
1. LH and FSH are produced in the pituitary, and estradiol and progesterone are produced in the ovaries.
2. Estradiol and progesterone secreted from the corpus luteum cause the endometrium to thicken.
3. Both progesterone and estradiol are produced by the follicles.
4. Secretion of GnRH by the hypothalamus is inhibited by low levels of estradiol but stimulated by high levels of estradiol.
Just prior to the middle of the cycle (approximately day 14), the high level of estrogen causes FSH and especially LH to rise rapidly, then fall. The spike in LH causes ovulation: the most mature follicle, like that shown in Figure \(3\), ruptures and releases its egg. The follicles that did not rupture degenerate and their eggs are lost. The level of estrogen decreases when the extra follicles degenerate.
Following ovulation, the ovarian cycle enters its luteal phase, illustrated in Figure \(4\) and the menstrual cycle enters its secretory phase, both of which run from about day 15 to 28. The luteal and secretory phases refer to changes in the ruptured follicle. The cells in the follicle undergo physical changes and produce a structure called a corpus luteum. The corpus luteum produces estrogen and progesterone. The progesterone facilitates the regrowth of the uterine lining and inhibits the release of further FSH and LH. The uterus is being prepared to accept a fertilized egg, should it occur during this cycle. The inhibition of FSH and LH prevents any further eggs and follicles from developing, while the progesterone is elevated. The level of estrogen produced by the corpus luteum increases to a steady level for the next few days.
If no fertilized egg is implanted into the uterus, the corpus luteum degenerates and the levels of estrogen and progesterone decrease. The endometrium begins to degenerate as the progesterone levels drop, initiating the next menstrual cycle. The decrease in progesterone also allows the hypothalamus to send GnRH to the anterior pituitary, releasing FSH and LH and starting the cycles again. Figure \(4\) visually compares the ovarian and uterine cycles as well as the commensurate hormone levels.
Art Connection
Which of the following statements about the menstrual cycle is false?
1. Progesterone levels rise during the luteal phase of the ovarian cycle and the secretory phase of the uterine cycle.
2. Menstruation occurs just after LH and FSH levels peak.
3. Menstruation occurs after progesterone levels drop.
4. Estrogen levels rise before ovulation, while progesterone levels rise after.
Menopause
As women approach their mid-40s to mid-50s, their ovaries begin to lose their sensitivity to FSH and LH. Menstrual periods become less frequent and finally cease; this is menopause. There are still eggs and potential follicles on the ovaries, but without the stimulation of FSH and LH, they will not produce a viable egg to be released. The outcome of this is the inability to have children.
The side effects of menopause include hot flashes, heavy sweating (especially at night), headaches, some hair loss, muscle pain, vaginal dryness, insomnia, depression, weight gain, and mood swings. Estrogen is involved in calcium metabolism and, without it, blood levels of calcium decrease. To replenish the blood, calcium is lost from bone which may decrease the bone density and lead to osteoporosis. Supplementation of estrogen in the form of hormone replacement therapy (HRT) can prevent bone loss, but the therapy can have negative side effects. While HRT is thought to give some protection from colon cancer, osteoporosis, heart disease, macular degeneration, and possibly depression, its negative side effects include increased risk of: stroke or heart attack, blood clots, breast cancer, ovarian cancer, endometrial cancer, gall bladder disease, and possibly dementia.
Career Connection: Reproductive Endocrinologist
A reproductive endocrinologist is a physician who treats a variety of hormonal disorders related to reproduction and infertility in both men and women. The disorders include menstrual problems, infertility, pregnancy loss, sexual dysfunction, and menopause. Doctors may use fertility drugs, surgery, or assisted reproductive techniques (ART) in their therapy. ART involves the use of procedures to manipulate the egg or sperm to facilitate reproduction, such as in vitro fertilization.
Reproductive endocrinologists undergo extensive medical training, first in a four-year residency in obstetrics and gynecology, then in a three-year fellowship in reproductive endocrinology. To be board certified in this area, the physician must pass written and oral exams in both areas.
Summary
The male and female reproductive cycles are controlled by hormones released from the hypothalamus and anterior pituitary as well as hormones from reproductive tissues and organs. The hypothalamus monitors the need for the FSH and LH hormones made and released from the anterior pituitary. FSH and LH affect reproductive structures to cause the formation of sperm and the preparation of eggs for release and possible fertilization. In the male, FSH and LH stimulate Sertoli cells and interstitial cells of Leydig in the testes to facilitate sperm production. The Leydig cells produce testosterone, which also is responsible for the secondary sexual characteristics of males. In females, FSH and LH cause estrogen and progesterone to be produced. They regulate the female reproductive system which is divided into the ovarian cycle and the menstrual cycle. Menopause occurs when the ovaries lose their sensitivity to FSH and LH and the female reproductive cycles slow to a stop.
Art Connections
Figure \(2\): Which of the following statements about hormone regulation of the female reproductive cycle is false?
1. LH and FSH are produced in the pituitary, and estradiol and progesterone are produced in the ovaries.
2. Estradiol and progesterone secreted from the corpus luteum cause the endometrium to thicken.
3. Both progesterone and estradiol are produced by the follicles.
4. Secretion of GnRH by the hypothalamus is inhibited by low levels of estradiol but stimulated by high levels of estradiol.
Answer
C
Figure \(4\): Which of the following statements about the menstrual cycle is false?
1. Progesterone levels rise during the luteal phase of the ovarian cycle and the secretory phase of the uterine cycle.
2. Menstruation occurs just after LH and FSH levels peak.
3. Menstruation occurs after progesterone levels drop.
4. Estrogen levels rise before ovulation, while progesterone levels rise after.
Answer
B
Glossary
estrogen
reproductive hormone in females that assists in endometrial regrowth, ovulation, and calcium absorption
follicle stimulating hormone (FSH)
reproductive hormone that causes sperm production in men and follicle development in women
gonadotropin-releasing hormone (GnRH)
hormone from the hypothalamus that causes the release of FSH and LH from the anterior pituitary
inhibin
hormone made by Sertoli cells; provides negative feedback to hypothalamus in control of FSH and GnRH release
interstitial cell of Leydig
cell in seminiferous tubules that makes testosterone
luteinizing hormone (LH)
reproductive hormone in both men and women, causes testosterone production in men and ovulation and lactation in women
menopause
loss of reproductive capacity in women due to decreased sensitivity of the ovaries to FSH and LH
menstrual cycle
cycle of the degradation and re-growth of the endometrium
ovarian cycle
cycle of preparation of egg for ovulation and the conversion of the follicle to the corpus luteum
ovulation
release of the egg by the most mature follicle
progesterone
reproductive hormone in women; assists in endometrial re-growth and inhibition of FSH and LH release
Sertoli cell
cell in seminiferous tubules that assists developing sperm and makes inhibin
testosterone
reproductive hormone in men that assists in sperm production and promoting secondary sexual characteristics | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/51%3A_The_Reproductive_System/51.04%3A_Structure_and_Function_of_the_Female_Reproductive_System/51.4.01%3A_Hormonal_Control_of_Human_Reproduction.txt |
Skills to Develop
• Explain fetal development during the three trimesters of gestation
• Describe labor and delivery
• Compare the efficacy and duration of various types of contraception
• Discuss causes of infertility and the therapeutic options available
Pregnancy begins with the fertilization of an egg and continues through to the birth of the individual. The length of time of gestation varies among animals, but is very similar among the great apes: human gestation is 266 days, while chimpanzee gestation is 237 days, a gorilla’s is 257 days, and orangutan gestation is 260 days long. The fox has a 57-day gestation. Dogs and cats have similar gestations averaging 60 days. The longest gestation for a land mammal is an African elephant at 640 days. The longest gestations among marine mammals are the beluga and sperm whales at 460 days.
Human Gestation
Twenty-four hours before fertilization, the egg has finished meiosis and becomes a mature oocyte. When fertilized (at conception) the egg becomes known as a zygote. The zygote travels through the oviduct to the uterus (Figure \(1\)). The developing embryo must implant into the wall of the uterus within seven days, or it will deteriorate and die. The outer layers of the zygote (blastocyst) grow into the endometrium by digesting the endometrial cells, and wound healing of the endometrium closes up the blastocyst into the tissue. Another layer of the blastocyst, the chorion, begins releasing a hormone called human beta chorionic gonadotropin (β-HCG) which makes its way to the corpus luteum and keeps that structure active. This ensures adequate levels of progesterone that will maintain the endometrium of the uterus for the support of the developing embryo. Pregnancy tests determine the level of β-HCG in urine or serum. If the hormone is present, the test is positive.
The gestation period is divided into three equal periods or trimesters. During the first two to four weeks of the first trimester, nutrition and waste are handled by the endometrial lining through diffusion. As the trimester progresses, the outer layer of the embryo begins to merge with the endometrium, and the placenta forms. This organ takes over the nutrient and waste requirements of the embryo and fetus, with the mother’s blood passing nutrients to the placenta and removing waste from it. Chemicals from the fetus, such as bilirubin, are processed by the mother’s liver for elimination. Some of the mother’s immunoglobulins will pass through the placenta, providing passive immunity against some potential infections.
Internal organs and body structures begin to develop during the first trimester. By five weeks, limb buds, eyes, the heart, and liver have been basically formed. By eight weeks, the term fetus applies, and the body is essentially formed, as shown in Figure \(2\). The individual is about five centimeters (two inches) in length and many of the organs, such as the lungs and liver, are not yet functioning. Exposure to any toxins is especially dangerous during the first trimester, as all of the body’s organs and structures are going through initial development. Anything that affects that development can have a severe effect on the fetus’ survival.
During the second trimester, the fetus grows to about 30 cm (12 inches), as shown in Figure \(3\). It becomes active and the mother usually feels the first movements. All organs and structures continue to develop. The placenta has taken over the functions of nutrition and waste and the production of estrogen and progesterone from the corpus luteum, which has degenerated. The placenta will continue functioning up through the delivery of the baby.
During the third trimester, the fetus grows to 3 to 4 kg (6 ½ -8 ½ lbs.) and about 50 cm (19-20 inches) long, as illustrated in Figure \(4\). This is the period of the most rapid growth during the pregnancy. Organ development continues to birth (and some systems, such as the nervous system and liver, continue to develop after birth). The mother will be at her most uncomfortable during this trimester. She may urinate frequently due to pressure on the bladder from the fetus. There may also be intestinal blockage and circulatory problems, especially in her legs. Clots may form in her legs due to pressure from the fetus on returning veins as they enter the abdominal cavity.
Link to Learning
Visit this site to see the stages of human fetal development.
Labor and Birth
Labor is the physical efforts of expulsion of the fetus and the placenta from the uterus during birth (parturition). Toward the end of the third trimester, estrogen causes receptors on the uterine wall to develop and bind the hormone oxytocin. At this time, the baby reorients, facing forward and down with the back or crown of the head engaging the cervix (uterine opening). This causes the cervix to stretch and nerve impulses are sent to the hypothalamus, which signals for the release of oxytocin from the posterior pituitary. The oxytocin causes the smooth muscle in the uterine wall to contract. At the same time, the placenta releases prostaglandins into the uterus, increasing the contractions. A positive feedback relay occurs between the uterus, hypothalamus, and the posterior pituitary to assure an adequate supply of oxytocin. As more smooth muscle cells are recruited, the contractions increase in intensity and force.
There are three stages to labor. During stage one, the cervix thins and dilates. This is necessary for the baby and placenta to be expelled during birth. The cervix will eventually dilate to about 10 cm. During stage two, the baby is expelled from the uterus. The uterus contracts and the mother pushes as she compresses her abdominal muscles to aid the delivery. The last stage is the passage of the placenta after the baby has been born and the organ has completely disengaged from the uterine wall. If labor should stop before stage two is reached, synthetic oxytocin, known as Pitocin, can be administered to restart and maintain labor.
An alternative to labor and delivery is the surgical delivery of the baby through a procedure called a Caesarian section. This is major abdominal surgery and can lead to post-surgical complications for the mother, but in some cases it may be the only way to safely deliver the baby.
The mother’s mammary glands go through changes during the third trimester to prepare for lactation and breastfeeding. When the baby begins suckling at the breast, signals are sent to the hypothalamus causing the release of prolactin from the anterior pituitary. Prolactin causes the mammary glands to produce milk. Oxytocin is also released, promoting the release of the milk. The milk contains nutrients for the baby’s development and growth as well as immunoglobulins to protect the child from bacterial and viral infections.
Contraception and Birth Control
The prevention of a pregnancy comes under the terms contraception or birth control. Strictly speaking, contraception refers to preventing the sperm and egg from joining. Both terms are, however, frequently used interchangeably.
Table \(1\): Contraceptive Methods
Method Examples Failure Rate in Typical Use Over 12 Months
Barrier male condom, female condom, sponge, cervical cap, diaphragm, spermicides 15 to 24%
Hormonal oral, patch, vaginal ring 8%
injection 3%
implant less than 1%
Other natural family planning 12 to 25%
withdrawal 27%
sterilization less than 1%
Table \(1\) lists common methods of contraception. The failure rates listed are not the ideal rates that could be realized, but the typical rates that occur. A failure rate is the number of pregnancies resulting from the method’s use over a twelve-month period. Barrier methods, such as condoms, cervical caps, and diaphragms, block sperm from entering the uterus, preventing fertilization. Spermicides are chemicals that are placed in the vagina that kill sperm. Sponges, which are saturated with spermicides, are placed in the vagina at the cervical opening. Combinations of spermicidal chemicals and barrier methods achieve lower failure rates than do the methods when used separately.
Nearly a quarter of the couples using barrier methods, natural family planning, or withdrawal can expect a failure of the method. Natural family planning is based on the monitoring of the menstrual cycle and having intercourse only during times when the egg is not available. A woman’s body temperature may rise a degree Celsius at ovulation and the cervical mucus may increase in volume and become more pliable. These changes give a general indication of when intercourse is more or less likely to result in fertilization. Withdrawal involves the removal of the penis from the vagina during intercourse, before ejaculation occurs. This is a risky method with a high failure rate due to the possible presence of sperm in the bulbourethral gland’s secretion, which may enter the vagina prior to removing the penis.
Hormonal methods use synthetic progesterone (sometimes in combination with estrogen), to inhibit the hypothalamus from releasing FSH or LH, and thus prevent an egg from being available for fertilization. The method of administering the hormone affects failure rate. The most reliable method, with a failure rate of less than 1 percent, is the implantation of the hormone under the skin. The same rate can be achieved through the sterilization procedures of vasectomy in the man or of tubal ligation in the woman, or by using an intrauterine device (IUD). IUDs are inserted into the uterus and establish an inflammatory condition that prevents fertilized eggs from implanting into the uterine wall.
Compliance with the contraceptive method is a strong contributor to the success or failure rate of any particular method. The only method that is completely effective at preventing conception is abstinence. The choice of contraceptive method depends on the goals of the woman or couple. Tubal ligation and vasectomy are considered permanent prevention, while other methods are reversible and provide short-term contraception.
Termination of an existing pregnancy can be spontaneous or voluntary. Spontaneous termination is a miscarriage and usually occurs very early in the pregnancy, usually within the first few weeks. This occurs when the fetus cannot develop properly and the gestation is naturally terminated. Voluntary termination of a pregnancy is an abortion. Laws regulating abortion vary between states and tend to view fetal viability as the criteria for allowing or preventing the procedure.
Infertility
Infertility is the inability to conceive a child or carry a child to birth. About 75 percent of causes of infertility can be identified; these include diseases, such as sexually transmitted diseases that can cause scarring of the reproductive tubes in either men or women, or developmental problems frequently related to abnormal hormone levels in one of the individuals. Inadequate nutrition, especially starvation, can delay menstruation. Stress can also lead to infertility. Short-term stress can affect hormone levels, while long-term stress can delay puberty and cause less frequent menstrual cycles. Other factors that affect fertility include toxins (such as cadmium), tobacco smoking, marijuana use, gonadal injuries, and aging.
If infertility is identified, several assisted reproductive technologies (ART) are available to aid conception. A common type of ART is in vitro fertilization (IVF) where an egg and sperm are combined outside the body and then placed in the uterus. Eggs are obtained from the woman after extensive hormonal treatments that prepare mature eggs for fertilization and prepare the uterus for implantation of the fertilized egg. Sperm are obtained from the man and they are combined with the eggs and supported through several cell divisions to ensure viability of the zygotes. When the embryos have reached the eight-cell stage, one or more is implanted into the woman’s uterus. If fertilization is not accomplished by simple IVF, a procedure that injects the sperm into an egg can be used. This is called intracytoplasmic sperm injection (ICSI) and is shown in Figure \(5\). IVF procedures produce a surplus of fertilized eggs and embryos that can be frozen and stored for future use. The procedures can also result in multiple births.
Summary
Human pregnancy begins with fertilization of an egg and proceeds through the three trimesters of gestation. The labor process has three stages (contractions, delivery of the fetus, expulsion of the placenta), each propelled by hormones. The first trimester lays down the basic structures of the body, including the limb buds, heart, eyes, and the liver. The second trimester continues the development of all of the organs and systems. The third trimester exhibits the greatest growth of the fetus and culminates in labor and delivery. Prevention of a pregnancy can be accomplished through a variety of methods including barriers, hormones, or other means. Assisted reproductive technologies may help individuals who have infertility problems.
Glossary
contraception
(also, birth control) various means used to prevent pregnancy
gestation
length of time for fetal development to birth
human beta chorionic gonadotropin (β-HCG)
hormone produced by the chorion of the zygote that helps to maintain the corpus luteum and elevated levels of progesterone
infertility
inability to conceive, carry, and deliver children
morning sickness
condition in the mother during the first trimester; includes feelings of nausea
placenta
organ that supports the diffusion of nutrients and waste between the mother’s and fetus’ blood | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/51%3A_The_Reproductive_System/51.04%3A_Structure_and_Function_of_the_Female_Reproductive_System/51.4.02%3A_Human_Pregnancy_and_Birth.txt |
Skills to Develop
• Explain fetal development during the three trimesters of gestation
• Describe labor and delivery
• Compare the efficacy and duration of various types of contraception
• Discuss causes of infertility and the therapeutic options available
Pregnancy begins with the fertilization of an egg and continues through to the birth of the individual. The length of time of gestation varies among animals, but is very similar among the great apes: human gestation is 266 days, while chimpanzee gestation is 237 days, a gorilla’s is 257 days, and orangutan gestation is 260 days long. The fox has a 57-day gestation. Dogs and cats have similar gestations averaging 60 days. The longest gestation for a land mammal is an African elephant at 640 days. The longest gestations among marine mammals are the beluga and sperm whales at 460 days.
Human Gestation
Twenty-four hours before fertilization, the egg has finished meiosis and becomes a mature oocyte. When fertilized (at conception) the egg becomes known as a zygote. The zygote travels through the oviduct to the uterus (Figure \(1\)). The developing embryo must implant into the wall of the uterus within seven days, or it will deteriorate and die. The outer layers of the zygote (blastocyst) grow into the endometrium by digesting the endometrial cells, and wound healing of the endometrium closes up the blastocyst into the tissue. Another layer of the blastocyst, the chorion, begins releasing a hormone called human beta chorionic gonadotropin (β-HCG) which makes its way to the corpus luteum and keeps that structure active. This ensures adequate levels of progesterone that will maintain the endometrium of the uterus for the support of the developing embryo. Pregnancy tests determine the level of β-HCG in urine or serum. If the hormone is present, the test is positive.
The gestation period is divided into three equal periods or trimesters. During the first two to four weeks of the first trimester, nutrition and waste are handled by the endometrial lining through diffusion. As the trimester progresses, the outer layer of the embryo begins to merge with the endometrium, and the placenta forms. This organ takes over the nutrient and waste requirements of the embryo and fetus, with the mother’s blood passing nutrients to the placenta and removing waste from it. Chemicals from the fetus, such as bilirubin, are processed by the mother’s liver for elimination. Some of the mother’s immunoglobulins will pass through the placenta, providing passive immunity against some potential infections.
Internal organs and body structures begin to develop during the first trimester. By five weeks, limb buds, eyes, the heart, and liver have been basically formed. By eight weeks, the term fetus applies, and the body is essentially formed, as shown in Figure \(2\). The individual is about five centimeters (two inches) in length and many of the organs, such as the lungs and liver, are not yet functioning. Exposure to any toxins is especially dangerous during the first trimester, as all of the body’s organs and structures are going through initial development. Anything that affects that development can have a severe effect on the fetus’ survival.
During the second trimester, the fetus grows to about 30 cm (12 inches), as shown in Figure \(3\). It becomes active and the mother usually feels the first movements. All organs and structures continue to develop. The placenta has taken over the functions of nutrition and waste and the production of estrogen and progesterone from the corpus luteum, which has degenerated. The placenta will continue functioning up through the delivery of the baby.
During the third trimester, the fetus grows to 3 to 4 kg (6 ½ -8 ½ lbs.) and about 50 cm (19-20 inches) long, as illustrated in Figure \(4\). This is the period of the most rapid growth during the pregnancy. Organ development continues to birth (and some systems, such as the nervous system and liver, continue to develop after birth). The mother will be at her most uncomfortable during this trimester. She may urinate frequently due to pressure on the bladder from the fetus. There may also be intestinal blockage and circulatory problems, especially in her legs. Clots may form in her legs due to pressure from the fetus on returning veins as they enter the abdominal cavity.
Link to Learning
Visit this site to see the stages of human fetal development.
Labor and Birth
Labor is the physical efforts of expulsion of the fetus and the placenta from the uterus during birth (parturition). Toward the end of the third trimester, estrogen causes receptors on the uterine wall to develop and bind the hormone oxytocin. At this time, the baby reorients, facing forward and down with the back or crown of the head engaging the cervix (uterine opening). This causes the cervix to stretch and nerve impulses are sent to the hypothalamus, which signals for the release of oxytocin from the posterior pituitary. The oxytocin causes the smooth muscle in the uterine wall to contract. At the same time, the placenta releases prostaglandins into the uterus, increasing the contractions. A positive feedback relay occurs between the uterus, hypothalamus, and the posterior pituitary to assure an adequate supply of oxytocin. As more smooth muscle cells are recruited, the contractions increase in intensity and force.
There are three stages to labor. During stage one, the cervix thins and dilates. This is necessary for the baby and placenta to be expelled during birth. The cervix will eventually dilate to about 10 cm. During stage two, the baby is expelled from the uterus. The uterus contracts and the mother pushes as she compresses her abdominal muscles to aid the delivery. The last stage is the passage of the placenta after the baby has been born and the organ has completely disengaged from the uterine wall. If labor should stop before stage two is reached, synthetic oxytocin, known as Pitocin, can be administered to restart and maintain labor.
An alternative to labor and delivery is the surgical delivery of the baby through a procedure called a Caesarian section. This is major abdominal surgery and can lead to post-surgical complications for the mother, but in some cases it may be the only way to safely deliver the baby.
The mother’s mammary glands go through changes during the third trimester to prepare for lactation and breastfeeding. When the baby begins suckling at the breast, signals are sent to the hypothalamus causing the release of prolactin from the anterior pituitary. Prolactin causes the mammary glands to produce milk. Oxytocin is also released, promoting the release of the milk. The milk contains nutrients for the baby’s development and growth as well as immunoglobulins to protect the child from bacterial and viral infections.
Contraception and Birth Control
The prevention of a pregnancy comes under the terms contraception or birth control. Strictly speaking, contraception refers to preventing the sperm and egg from joining. Both terms are, however, frequently used interchangeably.
Table \(1\): Contraceptive Methods
Method Examples Failure Rate in Typical Use Over 12 Months
Barrier male condom, female condom, sponge, cervical cap, diaphragm, spermicides 15 to 24%
Hormonal oral, patch, vaginal ring 8%
injection 3%
implant less than 1%
Other natural family planning 12 to 25%
withdrawal 27%
sterilization less than 1%
Table \(1\) lists common methods of contraception. The failure rates listed are not the ideal rates that could be realized, but the typical rates that occur. A failure rate is the number of pregnancies resulting from the method’s use over a twelve-month period. Barrier methods, such as condoms, cervical caps, and diaphragms, block sperm from entering the uterus, preventing fertilization. Spermicides are chemicals that are placed in the vagina that kill sperm. Sponges, which are saturated with spermicides, are placed in the vagina at the cervical opening. Combinations of spermicidal chemicals and barrier methods achieve lower failure rates than do the methods when used separately.
Nearly a quarter of the couples using barrier methods, natural family planning, or withdrawal can expect a failure of the method. Natural family planning is based on the monitoring of the menstrual cycle and having intercourse only during times when the egg is not available. A woman’s body temperature may rise a degree Celsius at ovulation and the cervical mucus may increase in volume and become more pliable. These changes give a general indication of when intercourse is more or less likely to result in fertilization. Withdrawal involves the removal of the penis from the vagina during intercourse, before ejaculation occurs. This is a risky method with a high failure rate due to the possible presence of sperm in the bulbourethral gland’s secretion, which may enter the vagina prior to removing the penis.
Hormonal methods use synthetic progesterone (sometimes in combination with estrogen), to inhibit the hypothalamus from releasing FSH or LH, and thus prevent an egg from being available for fertilization. The method of administering the hormone affects failure rate. The most reliable method, with a failure rate of less than 1 percent, is the implantation of the hormone under the skin. The same rate can be achieved through the sterilization procedures of vasectomy in the man or of tubal ligation in the woman, or by using an intrauterine device (IUD). IUDs are inserted into the uterus and establish an inflammatory condition that prevents fertilized eggs from implanting into the uterine wall.
Compliance with the contraceptive method is a strong contributor to the success or failure rate of any particular method. The only method that is completely effective at preventing conception is abstinence. The choice of contraceptive method depends on the goals of the woman or couple. Tubal ligation and vasectomy are considered permanent prevention, while other methods are reversible and provide short-term contraception.
Termination of an existing pregnancy can be spontaneous or voluntary. Spontaneous termination is a miscarriage and usually occurs very early in the pregnancy, usually within the first few weeks. This occurs when the fetus cannot develop properly and the gestation is naturally terminated. Voluntary termination of a pregnancy is an abortion. Laws regulating abortion vary between states and tend to view fetal viability as the criteria for allowing or preventing the procedure.
Infertility
Infertility is the inability to conceive a child or carry a child to birth. About 75 percent of causes of infertility can be identified; these include diseases, such as sexually transmitted diseases that can cause scarring of the reproductive tubes in either men or women, or developmental problems frequently related to abnormal hormone levels in one of the individuals. Inadequate nutrition, especially starvation, can delay menstruation. Stress can also lead to infertility. Short-term stress can affect hormone levels, while long-term stress can delay puberty and cause less frequent menstrual cycles. Other factors that affect fertility include toxins (such as cadmium), tobacco smoking, marijuana use, gonadal injuries, and aging.
If infertility is identified, several assisted reproductive technologies (ART) are available to aid conception. A common type of ART is in vitro fertilization (IVF) where an egg and sperm are combined outside the body and then placed in the uterus. Eggs are obtained from the woman after extensive hormonal treatments that prepare mature eggs for fertilization and prepare the uterus for implantation of the fertilized egg. Sperm are obtained from the man and they are combined with the eggs and supported through several cell divisions to ensure viability of the zygotes. When the embryos have reached the eight-cell stage, one or more is implanted into the woman’s uterus. If fertilization is not accomplished by simple IVF, a procedure that injects the sperm into an egg can be used. This is called intracytoplasmic sperm injection (ICSI) and is shown in Figure \(5\). IVF procedures produce a surplus of fertilized eggs and embryos that can be frozen and stored for future use. The procedures can also result in multiple births.
Summary
Human pregnancy begins with fertilization of an egg and proceeds through the three trimesters of gestation. The labor process has three stages (contractions, delivery of the fetus, expulsion of the placenta), each propelled by hormones. The first trimester lays down the basic structures of the body, including the limb buds, heart, eyes, and the liver. The second trimester continues the development of all of the organs and systems. The third trimester exhibits the greatest growth of the fetus and culminates in labor and delivery. Prevention of a pregnancy can be accomplished through a variety of methods including barriers, hormones, or other means. Assisted reproductive technologies may help individuals who have infertility problems.
Glossary
contraception
(also, birth control) various means used to prevent pregnancy
gestation
length of time for fetal development to birth
human beta chorionic gonadotropin (β-HCG)
hormone produced by the chorion of the zygote that helps to maintain the corpus luteum and elevated levels of progesterone
infertility
inability to conceive, carry, and deliver children
morning sickness
condition in the mother during the first trimester; includes feelings of nausea
placenta
organ that supports the diffusion of nutrients and waste between the mother’s and fetus’ blood | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/51%3A_The_Reproductive_System/51.05%3A_Contraception_and_Fertility_Treatments.txt |
Skills to Develop
• Discuss how fertilization occurs
• Explain how the embryo forms from the zygote
• Discuss the role of cleavage and gastrulation in animal development
The process in which an organism develops from a single-celled zygote to a multi-cellular organism is complex and well-regulated. The early stages of embryonic development are also crucial for ensuring the fitness of the organism.
Fertilization
Fertilization, pictured in Figure \(1\)a is the process in which gametes (an egg and sperm) fuse to form a zygote. The egg and sperm each contain one set of chromosomes. To ensure that the offspring has only one complete diploid set of chromosomes, only one sperm must fuse with one egg. In mammals, the egg is protected by a layer of extracellular matrix consisting mainly of glycoproteins called the zona pellucida. When a sperm binds to the zona pellucida, a series of biochemical events, called the acrosomal reactions, take place. In placental mammals, the acrosome contains digestive enzymes that initiate the degradation of the glycoprotein matrix protecting the egg and allowing the sperm plasma membrane to fuse with the egg plasma membrane, as illustrated in Figure \(1\)b. The fusion of these two membranes creates an opening through which the sperm nucleus is transferred into the ovum. The nuclear membranes of the egg and sperm break down and the two haploid genomes condense to form a diploid genome.
To ensure that no more than one sperm fertilizes the egg, once the acrosomal reactions take place at one location of the egg membrane, the egg releases proteins in other locations to prevent other sperm from fusing with the egg. If this mechanism fails, multiple sperm can fuse with the egg, resulting in polyspermy. The resulting embryo is not genetically viable and dies within a few days.
Cleavage and Blastula Stage
The development of multi-cellular organisms begins from a single-celled zygote, which undergoes rapid cell division to form the blastula. The rapid, multiple rounds of cell division are termed cleavage. Cleavage is illustrated in (Figure \(2\)a). After the cleavage has produced over 100 cells, the embryo is called a blastula. The blastula is usually a spherical layer of cells (the blastoderm) surrounding a fluid-filled or yolk-filled cavity (the blastocoel). Mammals at this stage form a structure called the blastocyst, characterized by an inner cell mass that is distinct from the surrounding blastula, shown in Figure \(2\)b. During cleavage, the cells divide without an increase in mass; that is, one large single-celled zygote divides into multiple smaller cells. Each cell within the blastula is called a blastomere.
Cleavage can take place in two ways: holoblastic (total) cleavage or meroblastic (partial) cleavage. The type of cleavage depends on the amount of yolk in the eggs. In placental mammals (including humans) where nourishment is provided by the mother’s body, the eggs have a very small amount of yolk and undergo holoblastic cleavage. Other species, such as birds, with a lot of yolk in the egg to nourish the embryo during development, undergo meroblastic cleavage.
In mammals, the blastula forms the blastocyst in the next stage of development. Here the cells in the blastula arrange themselves in two layers: the inner cell mass, and an outer layer called the trophoblast. The inner cell mass is also known as the embryoblast and this mass of cells will go on to form the embryo. At this stage of development, illustrated in Figure \(3\) the inner cell mass consists of embryonic stem cells that will differentiate into the different cell types needed by the organism. The trophoblast will contribute to the placenta and nourish the embryo.
Link to Learning
Visit the Virtual Human Embryo project at the Endowment for Human Development site to step through an interactive that shows the stages of embryo development, including micrographs and rotating 3-D images.
Gastrulation
The typical blastula is a ball of cells. The next stage in embryonic development is the formation of the body plan. The cells in the blastula rearrange themselves spatially to form three layers of cells. This process is called gastrulation. During gastrulation, the blastula folds upon itself to form the three layers of cells. Each of these layers is called a germ layer and each germ layer differentiates into different organ systems.
The three germs layers, shown in Figure \(4\), are the endoderm, the ectoderm, and the mesoderm. The ectoderm gives rise to the nervous system and the epidermis. The mesoderm gives rise to the muscle cells and connective tissue in the body. The endoderm gives rise to columnar cells found in the digestive system and many internal organs.
Everyday Connection: Are Designer Babies in Our Future?
If you could prevent your child from getting a devastating genetic disease, would you do it? Would you select the sex of your child or select for their attractiveness, strength, or intelligence? How far would you go to maximize the possibility of resistance to disease? The genetic engineering of a human child, the production of "designer babies" with desirable phenotypic characteristics, was once a topic restricted to science fiction. This is the case no longer: science fiction is now overlapping into science fact. Many phenotypic choices for offspring are already available, with many more likely to be possible in the not too distant future. Which traits should be selected and how they should be selected are topics of much debate within the worldwide medical community. The ethical and moral line is not always clear or agreed upon, and some fear that modern reproductive technologies could lead to a new form of eugenics.
Eugenics is the use of information and technology from a variety of sources to improve the genetic makeup of the human race. The goal of creating genetically superior humans was quite prevalent (although controversial) in several countries during the early 20th century, but fell into disrepute when Nazi Germany developed an extensive eugenics program in the 1930's and 40's. As part of their program, the Nazis forcibly sterilized hundreds of thousands of the so-called "unfit" and killed tens of thousands of institutionally disabled people as part of a systematic program to develop a genetically superior race of Germans known as Aryans. Ever since, eugenic ideas have not been as publicly expressed, but there are still those who promote them.
Efforts have been made in the past to control traits in human children using donated sperm from men with desired traits. In fact, eugenicist Robert Klark Graham established a sperm bank in 1980 that included samples exclusively from donors with high IQs. The "genius" sperm bank failed to capture the public's imagination and the operation closed in 1999.
In more recent times, the procedure known as prenatal genetic diagnosis (PGD) has been developed. PGD involves the screening of human embryos as part of the process of in vitro fertilization, during which embryos are conceived and grown outside the mother's body for some period of time before they are implanted. The term PGD usually refers to both the diagnosis, selection, and the implantation of the selected embryos.
In the least controversial use of PGD, embryos are tested for the presence of alleles which cause genetic diseases such as sickle cell disease, muscular dystrophy, and hemophilia, in which a single disease-causing allele or pair of alleles has been identified. By excluding embryos containing these alleles from implantation into the mother, the disease is prevented, and the unused embryos are either donated to science or discarded. There are relatively few in the worldwide medical community that question the ethics of this type of procedure, which allows individuals scared to have children because of the alleles they carry to do so successfully. The major limitation to this procedure is its expense. Not usually covered by medical insurance and thus out of reach financially for most couples, only a very small percentage of all live births use such complicated methodologies. Yet, even in cases like these where the ethical issues may seem to be clear-cut, not everyone agrees with the morality of these types of procedures. For example, to those who take the position that human life begins at conception, the discarding of unused embryos, a necessary result of PGD, is unacceptable under any circumstances.
A murkier ethical situation is found in the selection of a child's sex, which is easily performed by PGD. Currently, countries such as Great Britain have banned the selection of a child's sex for reasons other than preventing sex-linked diseases. Other countries allow the procedure for "family balancing", based on the desire of some parents to have at least one child of each sex. Still others, including the United States, have taken a scattershot approach to regulating these practices, essentially leaving it to the individual practicing physician to decide which practices are acceptable and which are not.
Even murkier are rare instances of disabled parents, such as those with deafness or dwarfism, who select embryos via PGD to ensure that they share their disability. These parents usually cite many positive aspects of their disabilities and associated culture as reasons for their choice, which they see as their moral right. To others, to purposely cause a disability in a child violates the basic medical principle of Primum non nocere, "first, do no harm." This procedure, although not illegal in most countries, demonstrates the complexity of ethical issues associated with choosing genetic traits in offspring.
Where could this process lead? Will this technology become more affordable and how should it be used? With the ability of technology to progress rapidly and unpredictably, a lack of definitive guidelines for the use of reproductive technologies before they arise might make it difficult for legislators to keep pace once they are in fact realized, assuming the process needs any government regulation at all. Other bioethicists argue that we should only deal with technologies that exist now, and not in some uncertain future. They argue that these types of procedures will always be expensive and rare, so the fears of eugenics and "master" races are unfounded and overstated. The debate continues.
Summary
The early stages of embryonic development begin with fertilization. The process of fertilization is tightly controlled to ensure that only one sperm fuses with one egg. After fertilization, the zygote undergoes cleavage to form the blastula. The blastula, which in some species is a hollow ball of cells, undergoes a process called gastrulation, in which the three germ layers form. The ectoderm gives rise to the nervous system and the epidermal skin cells, the mesoderm gives rise to the muscle cells and connective tissue in the body, and the endoderm gives rise to columnar cells and internal organs.
Glossary
acrosomal reaction
series of biochemical reactions that the sperm uses to break through the zona pellucida
blastocyst
structure formed when cells in the mammalian blastula separate into an inner and outer layer
gastrulation
process in which the blastula folds over itself to form the three germ layers
holoblastic
complete cleavage; takes place in cells with a small amount of yolk
inner cell mass
inner layer of cells in the blastocyst
meroblastic
partial cleavage; takes place in cells with a large amount of yolk
polyspermy
condition in which one egg is fertilized by multiple sperm
trophoblast
outer layer of cells in the blastocyst
zona pellucida
protective layer of glycoproteins on the mammalian egg | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/52%3A_Animal_Development/52.01%3A_Fertilization.txt |
Skills to Develop
• Discuss how fertilization occurs
• Explain how the embryo forms from the zygote
• Discuss the role of cleavage and gastrulation in animal development
The process in which an organism develops from a single-celled zygote to a multi-cellular organism is complex and well-regulated. The early stages of embryonic development are also crucial for ensuring the fitness of the organism.
Fertilization
Fertilization, pictured in Figure \(1\)a is the process in which gametes (an egg and sperm) fuse to form a zygote. The egg and sperm each contain one set of chromosomes. To ensure that the offspring has only one complete diploid set of chromosomes, only one sperm must fuse with one egg. In mammals, the egg is protected by a layer of extracellular matrix consisting mainly of glycoproteins called the zona pellucida. When a sperm binds to the zona pellucida, a series of biochemical events, called the acrosomal reactions, take place. In placental mammals, the acrosome contains digestive enzymes that initiate the degradation of the glycoprotein matrix protecting the egg and allowing the sperm plasma membrane to fuse with the egg plasma membrane, as illustrated in Figure \(1\)b. The fusion of these two membranes creates an opening through which the sperm nucleus is transferred into the ovum. The nuclear membranes of the egg and sperm break down and the two haploid genomes condense to form a diploid genome.
To ensure that no more than one sperm fertilizes the egg, once the acrosomal reactions take place at one location of the egg membrane, the egg releases proteins in other locations to prevent other sperm from fusing with the egg. If this mechanism fails, multiple sperm can fuse with the egg, resulting in polyspermy. The resulting embryo is not genetically viable and dies within a few days.
Cleavage and Blastula Stage
The development of multi-cellular organisms begins from a single-celled zygote, which undergoes rapid cell division to form the blastula. The rapid, multiple rounds of cell division are termed cleavage. Cleavage is illustrated in (Figure \(2\)a). After the cleavage has produced over 100 cells, the embryo is called a blastula. The blastula is usually a spherical layer of cells (the blastoderm) surrounding a fluid-filled or yolk-filled cavity (the blastocoel). Mammals at this stage form a structure called the blastocyst, characterized by an inner cell mass that is distinct from the surrounding blastula, shown in Figure \(2\)b. During cleavage, the cells divide without an increase in mass; that is, one large single-celled zygote divides into multiple smaller cells. Each cell within the blastula is called a blastomere.
Cleavage can take place in two ways: holoblastic (total) cleavage or meroblastic (partial) cleavage. The type of cleavage depends on the amount of yolk in the eggs. In placental mammals (including humans) where nourishment is provided by the mother’s body, the eggs have a very small amount of yolk and undergo holoblastic cleavage. Other species, such as birds, with a lot of yolk in the egg to nourish the embryo during development, undergo meroblastic cleavage.
In mammals, the blastula forms the blastocyst in the next stage of development. Here the cells in the blastula arrange themselves in two layers: the inner cell mass, and an outer layer called the trophoblast. The inner cell mass is also known as the embryoblast and this mass of cells will go on to form the embryo. At this stage of development, illustrated in Figure \(3\) the inner cell mass consists of embryonic stem cells that will differentiate into the different cell types needed by the organism. The trophoblast will contribute to the placenta and nourish the embryo.
Link to Learning
Visit the Virtual Human Embryo project at the Endowment for Human Development site to step through an interactive that shows the stages of embryo development, including micrographs and rotating 3-D images.
Gastrulation
The typical blastula is a ball of cells. The next stage in embryonic development is the formation of the body plan. The cells in the blastula rearrange themselves spatially to form three layers of cells. This process is called gastrulation. During gastrulation, the blastula folds upon itself to form the three layers of cells. Each of these layers is called a germ layer and each germ layer differentiates into different organ systems.
The three germs layers, shown in Figure \(4\), are the endoderm, the ectoderm, and the mesoderm. The ectoderm gives rise to the nervous system and the epidermis. The mesoderm gives rise to the muscle cells and connective tissue in the body. The endoderm gives rise to columnar cells found in the digestive system and many internal organs.
Everyday Connection: Are Designer Babies in Our Future?
If you could prevent your child from getting a devastating genetic disease, would you do it? Would you select the sex of your child or select for their attractiveness, strength, or intelligence? How far would you go to maximize the possibility of resistance to disease? The genetic engineering of a human child, the production of "designer babies" with desirable phenotypic characteristics, was once a topic restricted to science fiction. This is the case no longer: science fiction is now overlapping into science fact. Many phenotypic choices for offspring are already available, with many more likely to be possible in the not too distant future. Which traits should be selected and how they should be selected are topics of much debate within the worldwide medical community. The ethical and moral line is not always clear or agreed upon, and some fear that modern reproductive technologies could lead to a new form of eugenics.
Eugenics is the use of information and technology from a variety of sources to improve the genetic makeup of the human race. The goal of creating genetically superior humans was quite prevalent (although controversial) in several countries during the early 20th century, but fell into disrepute when Nazi Germany developed an extensive eugenics program in the 1930's and 40's. As part of their program, the Nazis forcibly sterilized hundreds of thousands of the so-called "unfit" and killed tens of thousands of institutionally disabled people as part of a systematic program to develop a genetically superior race of Germans known as Aryans. Ever since, eugenic ideas have not been as publicly expressed, but there are still those who promote them.
Efforts have been made in the past to control traits in human children using donated sperm from men with desired traits. In fact, eugenicist Robert Klark Graham established a sperm bank in 1980 that included samples exclusively from donors with high IQs. The "genius" sperm bank failed to capture the public's imagination and the operation closed in 1999.
In more recent times, the procedure known as prenatal genetic diagnosis (PGD) has been developed. PGD involves the screening of human embryos as part of the process of in vitro fertilization, during which embryos are conceived and grown outside the mother's body for some period of time before they are implanted. The term PGD usually refers to both the diagnosis, selection, and the implantation of the selected embryos.
In the least controversial use of PGD, embryos are tested for the presence of alleles which cause genetic diseases such as sickle cell disease, muscular dystrophy, and hemophilia, in which a single disease-causing allele or pair of alleles has been identified. By excluding embryos containing these alleles from implantation into the mother, the disease is prevented, and the unused embryos are either donated to science or discarded. There are relatively few in the worldwide medical community that question the ethics of this type of procedure, which allows individuals scared to have children because of the alleles they carry to do so successfully. The major limitation to this procedure is its expense. Not usually covered by medical insurance and thus out of reach financially for most couples, only a very small percentage of all live births use such complicated methodologies. Yet, even in cases like these where the ethical issues may seem to be clear-cut, not everyone agrees with the morality of these types of procedures. For example, to those who take the position that human life begins at conception, the discarding of unused embryos, a necessary result of PGD, is unacceptable under any circumstances.
A murkier ethical situation is found in the selection of a child's sex, which is easily performed by PGD. Currently, countries such as Great Britain have banned the selection of a child's sex for reasons other than preventing sex-linked diseases. Other countries allow the procedure for "family balancing", based on the desire of some parents to have at least one child of each sex. Still others, including the United States, have taken a scattershot approach to regulating these practices, essentially leaving it to the individual practicing physician to decide which practices are acceptable and which are not.
Even murkier are rare instances of disabled parents, such as those with deafness or dwarfism, who select embryos via PGD to ensure that they share their disability. These parents usually cite many positive aspects of their disabilities and associated culture as reasons for their choice, which they see as their moral right. To others, to purposely cause a disability in a child violates the basic medical principle of Primum non nocere, "first, do no harm." This procedure, although not illegal in most countries, demonstrates the complexity of ethical issues associated with choosing genetic traits in offspring.
Where could this process lead? Will this technology become more affordable and how should it be used? With the ability of technology to progress rapidly and unpredictably, a lack of definitive guidelines for the use of reproductive technologies before they arise might make it difficult for legislators to keep pace once they are in fact realized, assuming the process needs any government regulation at all. Other bioethicists argue that we should only deal with technologies that exist now, and not in some uncertain future. They argue that these types of procedures will always be expensive and rare, so the fears of eugenics and "master" races are unfounded and overstated. The debate continues.
Summary
The early stages of embryonic development begin with fertilization. The process of fertilization is tightly controlled to ensure that only one sperm fuses with one egg. After fertilization, the zygote undergoes cleavage to form the blastula. The blastula, which in some species is a hollow ball of cells, undergoes a process called gastrulation, in which the three germ layers form. The ectoderm gives rise to the nervous system and the epidermal skin cells, the mesoderm gives rise to the muscle cells and connective tissue in the body, and the endoderm gives rise to columnar cells and internal organs.
Glossary
acrosomal reaction
series of biochemical reactions that the sperm uses to break through the zona pellucida
blastocyst
structure formed when cells in the mammalian blastula separate into an inner and outer layer
gastrulation
process in which the blastula folds over itself to form the three germ layers
holoblastic
complete cleavage; takes place in cells with a small amount of yolk
inner cell mass
inner layer of cells in the blastocyst
meroblastic
partial cleavage; takes place in cells with a large amount of yolk
polyspermy
condition in which one egg is fertilized by multiple sperm
trophoblast
outer layer of cells in the blastocyst
zona pellucida
protective layer of glycoproteins on the mammalian egg | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/52%3A_Animal_Development/52.02%3A_Cleavage_and_the_Blastula_Stage.txt |
Skills to Develop
• Discuss how fertilization occurs
• Explain how the embryo forms from the zygote
• Discuss the role of cleavage and gastrulation in animal development
The process in which an organism develops from a single-celled zygote to a multi-cellular organism is complex and well-regulated. The early stages of embryonic development are also crucial for ensuring the fitness of the organism.
Fertilization
Fertilization, pictured in Figure \(1\)a is the process in which gametes (an egg and sperm) fuse to form a zygote. The egg and sperm each contain one set of chromosomes. To ensure that the offspring has only one complete diploid set of chromosomes, only one sperm must fuse with one egg. In mammals, the egg is protected by a layer of extracellular matrix consisting mainly of glycoproteins called the zona pellucida. When a sperm binds to the zona pellucida, a series of biochemical events, called the acrosomal reactions, take place. In placental mammals, the acrosome contains digestive enzymes that initiate the degradation of the glycoprotein matrix protecting the egg and allowing the sperm plasma membrane to fuse with the egg plasma membrane, as illustrated in Figure \(1\)b. The fusion of these two membranes creates an opening through which the sperm nucleus is transferred into the ovum. The nuclear membranes of the egg and sperm break down and the two haploid genomes condense to form a diploid genome.
To ensure that no more than one sperm fertilizes the egg, once the acrosomal reactions take place at one location of the egg membrane, the egg releases proteins in other locations to prevent other sperm from fusing with the egg. If this mechanism fails, multiple sperm can fuse with the egg, resulting in polyspermy. The resulting embryo is not genetically viable and dies within a few days.
Cleavage and Blastula Stage
The development of multi-cellular organisms begins from a single-celled zygote, which undergoes rapid cell division to form the blastula. The rapid, multiple rounds of cell division are termed cleavage. Cleavage is illustrated in (Figure \(2\)a). After the cleavage has produced over 100 cells, the embryo is called a blastula. The blastula is usually a spherical layer of cells (the blastoderm) surrounding a fluid-filled or yolk-filled cavity (the blastocoel). Mammals at this stage form a structure called the blastocyst, characterized by an inner cell mass that is distinct from the surrounding blastula, shown in Figure \(2\)b. During cleavage, the cells divide without an increase in mass; that is, one large single-celled zygote divides into multiple smaller cells. Each cell within the blastula is called a blastomere.
Cleavage can take place in two ways: holoblastic (total) cleavage or meroblastic (partial) cleavage. The type of cleavage depends on the amount of yolk in the eggs. In placental mammals (including humans) where nourishment is provided by the mother’s body, the eggs have a very small amount of yolk and undergo holoblastic cleavage. Other species, such as birds, with a lot of yolk in the egg to nourish the embryo during development, undergo meroblastic cleavage.
In mammals, the blastula forms the blastocyst in the next stage of development. Here the cells in the blastula arrange themselves in two layers: the inner cell mass, and an outer layer called the trophoblast. The inner cell mass is also known as the embryoblast and this mass of cells will go on to form the embryo. At this stage of development, illustrated in Figure \(3\) the inner cell mass consists of embryonic stem cells that will differentiate into the different cell types needed by the organism. The trophoblast will contribute to the placenta and nourish the embryo.
Link to Learning
Visit the Virtual Human Embryo project at the Endowment for Human Development site to step through an interactive that shows the stages of embryo development, including micrographs and rotating 3-D images.
Gastrulation
The typical blastula is a ball of cells. The next stage in embryonic development is the formation of the body plan. The cells in the blastula rearrange themselves spatially to form three layers of cells. This process is called gastrulation. During gastrulation, the blastula folds upon itself to form the three layers of cells. Each of these layers is called a germ layer and each germ layer differentiates into different organ systems.
The three germs layers, shown in Figure \(4\), are the endoderm, the ectoderm, and the mesoderm. The ectoderm gives rise to the nervous system and the epidermis. The mesoderm gives rise to the muscle cells and connective tissue in the body. The endoderm gives rise to columnar cells found in the digestive system and many internal organs.
Everyday Connection: Are Designer Babies in Our Future?
If you could prevent your child from getting a devastating genetic disease, would you do it? Would you select the sex of your child or select for their attractiveness, strength, or intelligence? How far would you go to maximize the possibility of resistance to disease? The genetic engineering of a human child, the production of "designer babies" with desirable phenotypic characteristics, was once a topic restricted to science fiction. This is the case no longer: science fiction is now overlapping into science fact. Many phenotypic choices for offspring are already available, with many more likely to be possible in the not too distant future. Which traits should be selected and how they should be selected are topics of much debate within the worldwide medical community. The ethical and moral line is not always clear or agreed upon, and some fear that modern reproductive technologies could lead to a new form of eugenics.
Eugenics is the use of information and technology from a variety of sources to improve the genetic makeup of the human race. The goal of creating genetically superior humans was quite prevalent (although controversial) in several countries during the early 20th century, but fell into disrepute when Nazi Germany developed an extensive eugenics program in the 1930's and 40's. As part of their program, the Nazis forcibly sterilized hundreds of thousands of the so-called "unfit" and killed tens of thousands of institutionally disabled people as part of a systematic program to develop a genetically superior race of Germans known as Aryans. Ever since, eugenic ideas have not been as publicly expressed, but there are still those who promote them.
Efforts have been made in the past to control traits in human children using donated sperm from men with desired traits. In fact, eugenicist Robert Klark Graham established a sperm bank in 1980 that included samples exclusively from donors with high IQs. The "genius" sperm bank failed to capture the public's imagination and the operation closed in 1999.
In more recent times, the procedure known as prenatal genetic diagnosis (PGD) has been developed. PGD involves the screening of human embryos as part of the process of in vitro fertilization, during which embryos are conceived and grown outside the mother's body for some period of time before they are implanted. The term PGD usually refers to both the diagnosis, selection, and the implantation of the selected embryos.
In the least controversial use of PGD, embryos are tested for the presence of alleles which cause genetic diseases such as sickle cell disease, muscular dystrophy, and hemophilia, in which a single disease-causing allele or pair of alleles has been identified. By excluding embryos containing these alleles from implantation into the mother, the disease is prevented, and the unused embryos are either donated to science or discarded. There are relatively few in the worldwide medical community that question the ethics of this type of procedure, which allows individuals scared to have children because of the alleles they carry to do so successfully. The major limitation to this procedure is its expense. Not usually covered by medical insurance and thus out of reach financially for most couples, only a very small percentage of all live births use such complicated methodologies. Yet, even in cases like these where the ethical issues may seem to be clear-cut, not everyone agrees with the morality of these types of procedures. For example, to those who take the position that human life begins at conception, the discarding of unused embryos, a necessary result of PGD, is unacceptable under any circumstances.
A murkier ethical situation is found in the selection of a child's sex, which is easily performed by PGD. Currently, countries such as Great Britain have banned the selection of a child's sex for reasons other than preventing sex-linked diseases. Other countries allow the procedure for "family balancing", based on the desire of some parents to have at least one child of each sex. Still others, including the United States, have taken a scattershot approach to regulating these practices, essentially leaving it to the individual practicing physician to decide which practices are acceptable and which are not.
Even murkier are rare instances of disabled parents, such as those with deafness or dwarfism, who select embryos via PGD to ensure that they share their disability. These parents usually cite many positive aspects of their disabilities and associated culture as reasons for their choice, which they see as their moral right. To others, to purposely cause a disability in a child violates the basic medical principle of Primum non nocere, "first, do no harm." This procedure, although not illegal in most countries, demonstrates the complexity of ethical issues associated with choosing genetic traits in offspring.
Where could this process lead? Will this technology become more affordable and how should it be used? With the ability of technology to progress rapidly and unpredictably, a lack of definitive guidelines for the use of reproductive technologies before they arise might make it difficult for legislators to keep pace once they are in fact realized, assuming the process needs any government regulation at all. Other bioethicists argue that we should only deal with technologies that exist now, and not in some uncertain future. They argue that these types of procedures will always be expensive and rare, so the fears of eugenics and "master" races are unfounded and overstated. The debate continues.
Summary
The early stages of embryonic development begin with fertilization. The process of fertilization is tightly controlled to ensure that only one sperm fuses with one egg. After fertilization, the zygote undergoes cleavage to form the blastula. The blastula, which in some species is a hollow ball of cells, undergoes a process called gastrulation, in which the three germ layers form. The ectoderm gives rise to the nervous system and the epidermal skin cells, the mesoderm gives rise to the muscle cells and connective tissue in the body, and the endoderm gives rise to columnar cells and internal organs.
Glossary
acrosomal reaction
series of biochemical reactions that the sperm uses to break through the zona pellucida
blastocyst
structure formed when cells in the mammalian blastula separate into an inner and outer layer
gastrulation
process in which the blastula folds over itself to form the three germ layers
holoblastic
complete cleavage; takes place in cells with a small amount of yolk
inner cell mass
inner layer of cells in the blastocyst
meroblastic
partial cleavage; takes place in cells with a large amount of yolk
polyspermy
condition in which one egg is fertilized by multiple sperm
trophoblast
outer layer of cells in the blastocyst
zona pellucida
protective layer of glycoproteins on the mammalian egg | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/52%3A_Animal_Development/52.03%3A_Gastrulation.txt |
Skills to Develop
• Describe the process of organogenesis
• Identify the anatomical axes formed in vertebrates
Gastrulation leads to the formation of the three germ layers that give rise, during further development, to the different organs in the animal body. This process is called organogenesis. Organogenesis is characterized by rapid and precise movements of the cells within the embryo.
Organogenesis
Organs form from the germ layers through the process of differentiation. During differentiation, the embryonic stem cells express specific sets of genes which will determine their ultimate cell type. For example, some cells in the ectoderm will express the genes specific to skin cells. As a result, these cells will differentiate into epidermal cells. The process of differentiation is regulated by cellular signaling cascades.
Scientists study organogenesis extensively in the lab in fruit flies (Drosophila) and the nematode Caenorhabditis elegans. Drosophila have segments along their bodies, and the patterning associated with the segment formation has allowed scientists to study which genes play important roles in organogenesis along the length of the embryo at different time points. The nematode C.elegans has roughly 1000 somatic cells and scientists have studied the fate of each of these cells during their development in the nematode life cycle. There is little variation in patterns of cell lineage between individuals, unlike in mammals where cell development from the embryo is dependent on cellular cues.
In vertebrates, one of the primary steps during organogenesis is the formation of the neural system. The ectoderm forms epithelial cells and tissues, and neuronal tissues. During the formation of the neural system, special signaling molecules called growth factors signal some cells at the edge of the ectoderm to become epidermis cells. The remaining cells in the center form the neural plate. If the signaling by growth factors were disrupted, then the entire ectoderm would differentiate into neural tissue.
The neural plate undergoes a series of cell movements where it rolls up and forms a tube called the neural tube, as illustrated in Figure \(1\). In further development, the neural tube will give rise to the brain and the spinal cord.
The mesoderm that lies on either side of the vertebrate neural tube will develop into the various connective tissues of the animal body. A spatial pattern of gene expression reorganizes the mesoderm into groups of cells called somites with spaces between them. The somites, illustrated in Figure \(2\) will further develop into the ribs, lungs, and segmental (spine) muscle. The mesoderm also forms a structure called the notochord, which is rod-shaped and forms the central axis of the animal body.
Vertebrate Axis Formation
Even as the germ layers form, the ball of cells still retains its spherical shape. However, animal bodies have lateral-medial (left-right), dorsal-ventral (back-belly), and anterior-posterior (head-feet) axes, illustrated in Figure \(3\).
How are these established? In one of the most seminal experiments ever to be carried out in developmental biology, Spemann and Mangold took dorsal cells from one embryo and transplanted them into the belly region of another embryo. They found that the transplanted embryo now had two notochords: one at the dorsal site from the original cells and another at the transplanted site. This suggested that the dorsal cells were genetically programmed to form the notochord and define the axis. Since then, researchers have identified many genes that are responsible for axis formation. Mutations in these genes leads to the loss of symmetry required for organism development.
Animal bodies have externally visible symmetry. However, the internal organs are not symmetric. For example, the heart is on the left side and the liver on the right. The formation of the central left-right axis is an important process during development. This internal asymmetry is established very early during development and involves many genes. Research is still ongoing to fully understand the developmental implications of these genes.
Summary
Organogenesis is the formation of organs from the germ layers. Each germ layer gives rise to specific tissue types. The first stage is the formation of the neural system in the ectoderm. The mesoderm gives rise to somites and the notochord. Formation of vertebrate axis is another important developmental stage.
Glossary
neural tube
tube-like structure that forms from the ectoderm and gives rise to the brain and spinal cord
organogenesis
process of organ formation
somite
group of cells separated by small spaces that form from the mesoderm and give rise to connective tissue | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/52%3A_Animal_Development/52.04%3A_Organogenesis.txt |
Skills to Develop
• Describe the process of organogenesis
• Identify the anatomical axes formed in vertebrates
Gastrulation leads to the formation of the three germ layers that give rise, during further development, to the different organs in the animal body. This process is called organogenesis. Organogenesis is characterized by rapid and precise movements of the cells within the embryo.
Organogenesis
Organs form from the germ layers through the process of differentiation. During differentiation, the embryonic stem cells express specific sets of genes which will determine their ultimate cell type. For example, some cells in the ectoderm will express the genes specific to skin cells. As a result, these cells will differentiate into epidermal cells. The process of differentiation is regulated by cellular signaling cascades.
Scientists study organogenesis extensively in the lab in fruit flies (Drosophila) and the nematode Caenorhabditis elegans. Drosophila have segments along their bodies, and the patterning associated with the segment formation has allowed scientists to study which genes play important roles in organogenesis along the length of the embryo at different time points. The nematode C.elegans has roughly 1000 somatic cells and scientists have studied the fate of each of these cells during their development in the nematode life cycle. There is little variation in patterns of cell lineage between individuals, unlike in mammals where cell development from the embryo is dependent on cellular cues.
In vertebrates, one of the primary steps during organogenesis is the formation of the neural system. The ectoderm forms epithelial cells and tissues, and neuronal tissues. During the formation of the neural system, special signaling molecules called growth factors signal some cells at the edge of the ectoderm to become epidermis cells. The remaining cells in the center form the neural plate. If the signaling by growth factors were disrupted, then the entire ectoderm would differentiate into neural tissue.
The neural plate undergoes a series of cell movements where it rolls up and forms a tube called the neural tube, as illustrated in Figure \(1\). In further development, the neural tube will give rise to the brain and the spinal cord.
The mesoderm that lies on either side of the vertebrate neural tube will develop into the various connective tissues of the animal body. A spatial pattern of gene expression reorganizes the mesoderm into groups of cells called somites with spaces between them. The somites, illustrated in Figure \(2\) will further develop into the ribs, lungs, and segmental (spine) muscle. The mesoderm also forms a structure called the notochord, which is rod-shaped and forms the central axis of the animal body.
Vertebrate Axis Formation
Even as the germ layers form, the ball of cells still retains its spherical shape. However, animal bodies have lateral-medial (left-right), dorsal-ventral (back-belly), and anterior-posterior (head-feet) axes, illustrated in Figure \(3\).
How are these established? In one of the most seminal experiments ever to be carried out in developmental biology, Spemann and Mangold took dorsal cells from one embryo and transplanted them into the belly region of another embryo. They found that the transplanted embryo now had two notochords: one at the dorsal site from the original cells and another at the transplanted site. This suggested that the dorsal cells were genetically programmed to form the notochord and define the axis. Since then, researchers have identified many genes that are responsible for axis formation. Mutations in these genes leads to the loss of symmetry required for organism development.
Animal bodies have externally visible symmetry. However, the internal organs are not symmetric. For example, the heart is on the left side and the liver on the right. The formation of the central left-right axis is an important process during development. This internal asymmetry is established very early during development and involves many genes. Research is still ongoing to fully understand the developmental implications of these genes.
Summary
Organogenesis is the formation of organs from the germ layers. Each germ layer gives rise to specific tissue types. The first stage is the formation of the neural system in the ectoderm. The mesoderm gives rise to somites and the notochord. Formation of vertebrate axis is another important developmental stage.
Glossary
neural tube
tube-like structure that forms from the ectoderm and gives rise to the brain and spinal cord
organogenesis
process of organ formation
somite
group of cells separated by small spaces that form from the mesoderm and give rise to connective tissue | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/52%3A_Animal_Development/52.05%3A_Vertebrate_Axis_Pattern_Formation.txt |
Skills to Develop
• Explain fetal development during the three trimesters of gestation
• Describe labor and delivery
• Compare the efficacy and duration of various types of contraception
• Discuss causes of infertility and the therapeutic options available
Pregnancy begins with the fertilization of an egg and continues through to the birth of the individual. The length of time of gestation varies among animals, but is very similar among the great apes: human gestation is 266 days, while chimpanzee gestation is 237 days, a gorilla’s is 257 days, and orangutan gestation is 260 days long. The fox has a 57-day gestation. Dogs and cats have similar gestations averaging 60 days. The longest gestation for a land mammal is an African elephant at 640 days. The longest gestations among marine mammals are the beluga and sperm whales at 460 days.
Human Gestation
Twenty-four hours before fertilization, the egg has finished meiosis and becomes a mature oocyte. When fertilized (at conception) the egg becomes known as a zygote. The zygote travels through the oviduct to the uterus (Figure \(1\)). The developing embryo must implant into the wall of the uterus within seven days, or it will deteriorate and die. The outer layers of the zygote (blastocyst) grow into the endometrium by digesting the endometrial cells, and wound healing of the endometrium closes up the blastocyst into the tissue. Another layer of the blastocyst, the chorion, begins releasing a hormone called human beta chorionic gonadotropin (β-HCG) which makes its way to the corpus luteum and keeps that structure active. This ensures adequate levels of progesterone that will maintain the endometrium of the uterus for the support of the developing embryo. Pregnancy tests determine the level of β-HCG in urine or serum. If the hormone is present, the test is positive.
The gestation period is divided into three equal periods or trimesters. During the first two to four weeks of the first trimester, nutrition and waste are handled by the endometrial lining through diffusion. As the trimester progresses, the outer layer of the embryo begins to merge with the endometrium, and the placenta forms. This organ takes over the nutrient and waste requirements of the embryo and fetus, with the mother’s blood passing nutrients to the placenta and removing waste from it. Chemicals from the fetus, such as bilirubin, are processed by the mother’s liver for elimination. Some of the mother’s immunoglobulins will pass through the placenta, providing passive immunity against some potential infections.
Internal organs and body structures begin to develop during the first trimester. By five weeks, limb buds, eyes, the heart, and liver have been basically formed. By eight weeks, the term fetus applies, and the body is essentially formed, as shown in Figure \(2\). The individual is about five centimeters (two inches) in length and many of the organs, such as the lungs and liver, are not yet functioning. Exposure to any toxins is especially dangerous during the first trimester, as all of the body’s organs and structures are going through initial development. Anything that affects that development can have a severe effect on the fetus’ survival.
During the second trimester, the fetus grows to about 30 cm (12 inches), as shown in Figure \(3\). It becomes active and the mother usually feels the first movements. All organs and structures continue to develop. The placenta has taken over the functions of nutrition and waste and the production of estrogen and progesterone from the corpus luteum, which has degenerated. The placenta will continue functioning up through the delivery of the baby.
During the third trimester, the fetus grows to 3 to 4 kg (6 ½ -8 ½ lbs.) and about 50 cm (19-20 inches) long, as illustrated in Figure \(4\). This is the period of the most rapid growth during the pregnancy. Organ development continues to birth (and some systems, such as the nervous system and liver, continue to develop after birth). The mother will be at her most uncomfortable during this trimester. She may urinate frequently due to pressure on the bladder from the fetus. There may also be intestinal blockage and circulatory problems, especially in her legs. Clots may form in her legs due to pressure from the fetus on returning veins as they enter the abdominal cavity.
Link to Learning
Visit this site to see the stages of human fetal development.
Labor and Birth
Labor is the physical efforts of expulsion of the fetus and the placenta from the uterus during birth (parturition). Toward the end of the third trimester, estrogen causes receptors on the uterine wall to develop and bind the hormone oxytocin. At this time, the baby reorients, facing forward and down with the back or crown of the head engaging the cervix (uterine opening). This causes the cervix to stretch and nerve impulses are sent to the hypothalamus, which signals for the release of oxytocin from the posterior pituitary. The oxytocin causes the smooth muscle in the uterine wall to contract. At the same time, the placenta releases prostaglandins into the uterus, increasing the contractions. A positive feedback relay occurs between the uterus, hypothalamus, and the posterior pituitary to assure an adequate supply of oxytocin. As more smooth muscle cells are recruited, the contractions increase in intensity and force.
There are three stages to labor. During stage one, the cervix thins and dilates. This is necessary for the baby and placenta to be expelled during birth. The cervix will eventually dilate to about 10 cm. During stage two, the baby is expelled from the uterus. The uterus contracts and the mother pushes as she compresses her abdominal muscles to aid the delivery. The last stage is the passage of the placenta after the baby has been born and the organ has completely disengaged from the uterine wall. If labor should stop before stage two is reached, synthetic oxytocin, known as Pitocin, can be administered to restart and maintain labor.
An alternative to labor and delivery is the surgical delivery of the baby through a procedure called a Caesarian section. This is major abdominal surgery and can lead to post-surgical complications for the mother, but in some cases it may be the only way to safely deliver the baby.
The mother’s mammary glands go through changes during the third trimester to prepare for lactation and breastfeeding. When the baby begins suckling at the breast, signals are sent to the hypothalamus causing the release of prolactin from the anterior pituitary. Prolactin causes the mammary glands to produce milk. Oxytocin is also released, promoting the release of the milk. The milk contains nutrients for the baby’s development and growth as well as immunoglobulins to protect the child from bacterial and viral infections.
Contraception and Birth Control
The prevention of a pregnancy comes under the terms contraception or birth control. Strictly speaking, contraception refers to preventing the sperm and egg from joining. Both terms are, however, frequently used interchangeably.
Table \(1\): Contraceptive Methods
Method Examples Failure Rate in Typical Use Over 12 Months
Barrier male condom, female condom, sponge, cervical cap, diaphragm, spermicides 15 to 24%
Hormonal oral, patch, vaginal ring 8%
injection 3%
implant less than 1%
Other natural family planning 12 to 25%
withdrawal 27%
sterilization less than 1%
Table \(1\) lists common methods of contraception. The failure rates listed are not the ideal rates that could be realized, but the typical rates that occur. A failure rate is the number of pregnancies resulting from the method’s use over a twelve-month period. Barrier methods, such as condoms, cervical caps, and diaphragms, block sperm from entering the uterus, preventing fertilization. Spermicides are chemicals that are placed in the vagina that kill sperm. Sponges, which are saturated with spermicides, are placed in the vagina at the cervical opening. Combinations of spermicidal chemicals and barrier methods achieve lower failure rates than do the methods when used separately.
Nearly a quarter of the couples using barrier methods, natural family planning, or withdrawal can expect a failure of the method. Natural family planning is based on the monitoring of the menstrual cycle and having intercourse only during times when the egg is not available. A woman’s body temperature may rise a degree Celsius at ovulation and the cervical mucus may increase in volume and become more pliable. These changes give a general indication of when intercourse is more or less likely to result in fertilization. Withdrawal involves the removal of the penis from the vagina during intercourse, before ejaculation occurs. This is a risky method with a high failure rate due to the possible presence of sperm in the bulbourethral gland’s secretion, which may enter the vagina prior to removing the penis.
Hormonal methods use synthetic progesterone (sometimes in combination with estrogen), to inhibit the hypothalamus from releasing FSH or LH, and thus prevent an egg from being available for fertilization. The method of administering the hormone affects failure rate. The most reliable method, with a failure rate of less than 1 percent, is the implantation of the hormone under the skin. The same rate can be achieved through the sterilization procedures of vasectomy in the man or of tubal ligation in the woman, or by using an intrauterine device (IUD). IUDs are inserted into the uterus and establish an inflammatory condition that prevents fertilized eggs from implanting into the uterine wall.
Compliance with the contraceptive method is a strong contributor to the success or failure rate of any particular method. The only method that is completely effective at preventing conception is abstinence. The choice of contraceptive method depends on the goals of the woman or couple. Tubal ligation and vasectomy are considered permanent prevention, while other methods are reversible and provide short-term contraception.
Termination of an existing pregnancy can be spontaneous or voluntary. Spontaneous termination is a miscarriage and usually occurs very early in the pregnancy, usually within the first few weeks. This occurs when the fetus cannot develop properly and the gestation is naturally terminated. Voluntary termination of a pregnancy is an abortion. Laws regulating abortion vary between states and tend to view fetal viability as the criteria for allowing or preventing the procedure.
Infertility
Infertility is the inability to conceive a child or carry a child to birth. About 75 percent of causes of infertility can be identified; these include diseases, such as sexually transmitted diseases that can cause scarring of the reproductive tubes in either men or women, or developmental problems frequently related to abnormal hormone levels in one of the individuals. Inadequate nutrition, especially starvation, can delay menstruation. Stress can also lead to infertility. Short-term stress can affect hormone levels, while long-term stress can delay puberty and cause less frequent menstrual cycles. Other factors that affect fertility include toxins (such as cadmium), tobacco smoking, marijuana use, gonadal injuries, and aging.
If infertility is identified, several assisted reproductive technologies (ART) are available to aid conception. A common type of ART is in vitro fertilization (IVF) where an egg and sperm are combined outside the body and then placed in the uterus. Eggs are obtained from the woman after extensive hormonal treatments that prepare mature eggs for fertilization and prepare the uterus for implantation of the fertilized egg. Sperm are obtained from the man and they are combined with the eggs and supported through several cell divisions to ensure viability of the zygotes. When the embryos have reached the eight-cell stage, one or more is implanted into the woman’s uterus. If fertilization is not accomplished by simple IVF, a procedure that injects the sperm into an egg can be used. This is called intracytoplasmic sperm injection (ICSI) and is shown in Figure \(5\). IVF procedures produce a surplus of fertilized eggs and embryos that can be frozen and stored for future use. The procedures can also result in multiple births.
Summary
Human pregnancy begins with fertilization of an egg and proceeds through the three trimesters of gestation. The labor process has three stages (contractions, delivery of the fetus, expulsion of the placenta), each propelled by hormones. The first trimester lays down the basic structures of the body, including the limb buds, heart, eyes, and the liver. The second trimester continues the development of all of the organs and systems. The third trimester exhibits the greatest growth of the fetus and culminates in labor and delivery. Prevention of a pregnancy can be accomplished through a variety of methods including barriers, hormones, or other means. Assisted reproductive technologies may help individuals who have infertility problems.
Glossary
contraception
(also, birth control) various means used to prevent pregnancy
gestation
length of time for fetal development to birth
human beta chorionic gonadotropin (β-HCG)
hormone produced by the chorion of the zygote that helps to maintain the corpus luteum and elevated levels of progesterone
infertility
inability to conceive, carry, and deliver children
morning sickness
condition in the mother during the first trimester; includes feelings of nausea
placenta
organ that supports the diffusion of nutrients and waste between the mother’s and fetus’ blood | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/52%3A_Animal_Development/52.06%3A_Human_Development.txt |
Skills to Develop
• Compare innate and learned behavior
• Discuss how movement and migration behaviors are a result of natural selection
• Discuss the different ways members of a population communicate with each other
• Give examples of how species use energy for mating displays and other courtship behaviors
• Differentiate between various mating systems
• Describe different ways that species learn
Behavior is the change in activity of an organism in response to a stimulus. Behavioral biology is the study of the biological and evolutionary bases for such changes. The idea that behaviors evolved as a result of the pressures of natural selection is not new. Animal behavior has been studied for decades, by biologists in the science of ethology, by psychologists in the science of comparative psychology, and by scientists of many disciplines in the study of neurobiology. Although there is overlap between these disciplines, scientists in these behavioral fields take different approaches. Comparative psychology is an extension of work done in human and behavioral psychology. Ethology is an extension of genetics, evolution, anatomy, physiology, and other biological disciplines. Still, one cannot study behavioral biology without touching on both comparative psychology and ethology.
One goal of behavioral biology is to dissect out the innate behaviors, which have a strong genetic component and are largely independent of environmental influences, from the learned behaviors, which result from environmental conditioning. Innate behavior, or instinct, is important because there is no risk of an incorrect behavior being learned. They are “hard wired” into the system. On the other hand, learned behaviors, although riskier, are flexible, dynamic, and can be altered according to changes in the environment.
Innate Behaviors: Movement and Migration
Innate or instinctual behaviors rely on response to stimuli. The simplest example of this is a reflex action, an involuntary and rapid response to stimulus. To test the “knee-jerk” reflex, a doctor taps the patellar tendon below the kneecap with a rubber hammer. The stimulation of the nerves there leads to the reflex of extending the leg at the knee. This is similar to the reaction of someone who touches a hot stove and instinctually pulls his or her hand away. Even humans, with our great capacity to learn, still exhibit a variety of innate behaviors.
Kinesis and Taxis
Another activity or movement of innate behavior is kinesis, or the undirected movement in response to a stimulus. Orthokinesis is the increased or decreased speed of movement of an organism in response to a stimulus. Woodlice, for example, increase their speed of movement when exposed to high or low temperatures. This movement, although random, increases the probability that the insect spends less time in the unfavorable environment. Another example is klinokinesis, an increase in turning behaviors. It is exhibited by bacteria such as E. coli which, in association with orthokinesis, helps the organisms randomly find a more hospitable environment.
A similar, but more directed version of kinesis is taxis: the directed movement towards or away from a stimulus. This movement can be in response to light (phototaxis), chemical signals (chemotaxis), or gravity (geotaxis) and can be directed toward (positive) or away (negative) from the source of the stimulus. An example of a positive chemotaxis is exhibited by the unicellular protozoan Tetrahymena thermophila. This organism swims using its cilia, at times moving in a straight line, and at other times making turns. The attracting chemotactic agent alters the frequency of turning as the organism moves directly toward the source, following the increasing concentration gradient.
Fixed Action Patterns
A fixed action pattern is a series of movements elicited by a stimulus such that even when the stimulus is removed, the pattern goes on to completion. An example of such a behavior occurs in the three-spined stickleback, a small freshwater fish (Figure \(1\)). Males of this species develop a red belly during breeding season and show instinctual aggressiveness to other males during this time. In laboratory experiments, researchers exposed such fish to objects that in no way resemble a fish in their shape, but which were painted red on their lower halves. The male sticklebacks responded aggressively to the objects just as if they were real male sticklebacks.
Migration
Migration is the long-range seasonal movement of animals. It is an evolved, adapted response to variation in resource availability, and it is a common phenomenon found in all major groups of animals. Birds fly south for the winter to get to warmer climates with sufficient food, and salmon migrate to their spawning grounds. The popular 2005 documentary March of the Penguins followed the 62-mile migration of emperor penguins through Antarctica to bring food back to their breeding site and to their young. Wildebeests (Figure \(2\)) migrate over 1800 miles each year in search of new grasslands.
Although migration is thought of as innate behavior, only some migrating species always migrate (obligate migration). Animals that exhibit facultative migration can choose to migrate or not. Additionally, in some animals, only a portion of the population migrates, whereas the rest does not migrate (incomplete migration). For example, owls that live in the tundra may migrate in years when their food source, small rodents, is relatively scarce, but not migrate during the years when rodents are plentiful.
Foraging
Foraging is the act of searching for and exploiting food resources. Feeding behaviors that maximize energy gain and minimize energy expenditure are called optimal foraging behaviors, and these are favored by natural section. The painted stork, for example, uses its long beak to search the bottom of a freshwater marshland for crabs and other food (Figure \(3\)).
Innate Behaviors: Living in Groups
Not all animals live in groups, but even those that live relatively solitary lives, with the exception of those that can reproduce asexually, must mate. Mating usually involves one animal signaling another so as to communicate the desire to mate. There are several types of energy-intensive behaviors or displays associated with mating, called mating rituals. Other behaviors found in populations that live in groups are described in terms of which animal benefits from the behavior. In selfish behavior, only the animal in question benefits; in altruistic behavior, one animal’s actions benefit another animal; cooperative behavior describes when both animals benefit. All of these behaviors involve some sort of communication between population members.
Communication within a Species
Animals communicate with each other using stimuli known as signals. An example of this is seen in the three-spined stickleback, where the visual signal of a red region in the lower half of a fish signals males to become aggressive and signals females to mate. Other signals are chemical (pheromones), aural (sound), visual (courtship and aggressive displays), or tactile (touch). These types of communication may be instinctual or learned or a combination of both. These are not the same as the communication we associate with language, which has been observed only in humans and perhaps in some species of primates and cetaceans.
A pheromone is a secreted chemical signal used to obtain a response from another individual of the same species. The purpose of pheromones is to elicit a specific behavior from the receiving individual. Pheromones are especially common among social insects, but they are used by many species to attract the opposite sex, to sound alarms, to mark food trails, and to elicit other, more complex behaviors. Even humans are thought to respond to certain pheromones called axillary steroids. These chemicals influence human perception of other people, and in one study were responsible for a group of women synchronizing their menstrual cycles. The role of pheromones in human-to-human communication is still somewhat controversial and continues to be researched.
Songs are an example of an aural signal, one that needs to be heard by the recipient. Perhaps the best known of these are songs of birds, which identify the species and are used to attract mates. Other well-known songs are those of whales, which are of such low frequency that they can travel long distances underwater. Dolphins communicate with each other using a wide variety of vocalizations. Male crickets make chirping sounds using a specialized organ to attract a mate, repel other males, and to announce a successful mating.
Courtship displays are a series of ritualized visual behaviors (signals) designed to attract and convince a member of the opposite sex to mate. These displays are ubiquitous in the animal kingdom. Often these displays involve a series of steps, including an initial display by one member followed by a response from the other. If at any point, the display is performed incorrectly or a proper response is not given, the mating ritual is abandoned and the mating attempt will be unsuccessful. The mating display of the common stork is shown in Figure \(4\).
Aggressive displays are also common in the animal kingdom. An example is when a dog bares its teeth when it wants another dog to back down. Presumably, these displays communicate not only the willingness of the animal to fight, but also its fighting ability. Although these displays do signal aggression on the part of the sender, it is thought that these displays are actually a mechanism to reduce the amount of actual fighting that occurs between members of the same species: they allow individuals to assess the fighting ability of their opponent and thus decide whether it is “worth the fight.” The testing of certain hypotheses using game theory has led to the conclusion that some of these displays may overstate an animal’s actual fighting ability and are used to “bluff” the opponent. This type of interaction, even if “dishonest,” would be favored by natural selection if it is successful more times than not.
Distraction displays are seen in birds and some fish. They are designed to attract a predator away from the nest that contains their young. This is an example of an altruistic behavior: it benefits the young more than the individual performing the display, which is putting itself at risk by doing so.
Many animals, especially primates, communicate with other members in the group through touch. Activities such as grooming, touching the shoulder or root of the tail, embracing, lip contact, and greeting ceremonies have all been observed in the Indian langur, an Old World monkey. Similar behaviors are found in other primates, especially in the great apes.
Link to Learning
The killdeer bird distracts predators from its eggs by faking a broken wing display in this video taken in Boise, Idaho.
Altruistic Behaviors
Behaviors that lower the fitness of the individual but increase the fitness of another individual are termed altruistic. Examples of such behaviors are seen widely across the animal kingdom. Social insects such as worker bees have no ability to reproduce, yet they maintain the queen so she can populate the hive with her offspring. Meerkats keep a sentry standing guard to warn the rest of the colony about intruders, even though the sentry is putting itself at risk. Wolves and wild dogs bring meat to pack members not present during a hunt. Lemurs take care of infants unrelated to them. Although on the surface, these behaviors appear to be altruistic, it may not be so simple.
There has been much discussion over why altruistic behaviors exist. Do these behaviors lead to overall evolutionary advantages for their species? Do they help the altruistic individual pass on its own genes? And what about such activities between unrelated individuals? One explanation for altruistic-type behaviors is found in the genetics of natural selection. In the 1976 book, The Selfish Gene, scientist Richard Dawkins attempted to explain many seemingly altruistic behaviors from the viewpoint of the gene itself. Although a gene obviously cannot be selfish in the human sense, it may appear that way if the sacrifice of an individual benefits related individuals that share genes that are identical by descent (present in relatives because of common lineage). Mammal parents make this sacrifice to take care of their offspring. Emperor penguins migrate miles in harsh conditions to bring food back for their young. Selfish gene theory has been controversial over the years and is still discussed among scientists in related fields.
Even less-related individuals, those with less genetic identity than that shared by parent and offspring, benefit from seemingly altruistic behavior. The activities of social insects such as bees, wasps, ants, and termites are good examples. Sterile workers in these societies take care of the queen because they are closely related to it, and as the queen has offspring, she is passing on genes from the workers indirectly. Thus, it is of fitness benefit for the worker to maintain the queen without having any direct chance of passing on its genes due to its sterility. The lowering of individual fitness to enhance the reproductive fitness of a relative and thus one’s inclusive fitness evolves through kin selection. This phenomenon can explain many superficially altruistic behaviors seen in animals. However, these behaviors may not be truly defined as altruism in these cases because the actor is actually increasing its own fitness either directly (through its own offspring) or indirectly (through the inclusive fitness it gains through relatives that share genes with it).
Unrelated individuals may also act altruistically to each other, and this seems to defy the “selfish gene” explanation. An example of this observed in many monkey species where a monkey will present its back to an unrelated monkey to have that individual pick the parasites from its fur. After a certain amount of time, the roles are reversed and the first monkey now grooms the second monkey. Thus, there is reciprocity in the behavior. Both benefit from the interaction and their fitness is raised more than if neither cooperated nor if one cooperated and the other did not cooperate. This behavior is still not necessarily altruism, as the “giving” behavior of the actor is based on the expectation that it will be the “receiver” of the behavior in the future, termed reciprocal altruism. Reciprocal altruism requires that individuals repeatedly encounter each other, often the result of living in the same social group, and that cheaters (those that never “give back”) are punished.
Evolutionary game theory, a modification of classical game theory in mathematics, has shown that many of these so-called “altruistic behaviors” are not altruistic at all. The definition of “pure” altruism, based on human behavior, is an action that benefits another without any direct benefit to oneself. Most of the behaviors previously described do not seem to satisfy this definition, and game theorists are good at finding “selfish” components in them. Others have argued that the terms “selfish” and “altruistic” should be dropped completely when discussing animal behavior, as they describe human behavior and may not be directly applicable to instinctual animal activity. What is clear, though, is that heritable behaviors that improve the chances of passing on one’s genes or a portion of one’s genes are favored by natural selection and will be retained in future generations as long as those behaviors convey a fitness advantage. These instinctual behaviors may then be applied, in special circumstances, to other species, as long as it doesn’t lower the animal’s fitness.
Finding Sex Partners
Not all animals reproduce sexually, but many that do have the same challenge: they need to find a suitable mate and often have to compete with other individuals to obtain one. Significant energy is spent in the process of locating, attracting, and mating with the sex partner. Two types of selection occur during this process and can lead to traits that are important to reproduction called secondary sexual characteristics: intersexual selection, the choosing of a mate where individuals of one sex choose mates of the other sex, and intrasexual selection, the competition for mates between species members of the same sex. Intersexual selection is often complex because choosing a mate may be based on a variety of visual, aural, tactile, and chemical cues. An example of intersexual selection is when female peacocks choose to mate with the male with the brightest plumage. This type of selection often leads to traits in the chosen sex that do not enhance survival, but are those traits most attractive to the opposite sex (often at the expense of survival). Intrasexual selection involves mating displays and aggressive mating rituals such as rams butting heads—the winner of these battles is the one that is able to mate. Many of these rituals use up considerable energy but result in the selection of the healthiest, strongest, and/or most dominant individuals for mating. Three general mating systems, all involving innate as opposed to learned behaviors, are seen in animal populations: monogamous, polygynous, and polyandrous.
In monogamous systems, one male and one female are paired for at least one breeding season. In some animals, such as the gray wolf, these associations can last much longer, even a lifetime. Several explanations have been proposed for this type of mating system. The “mate-guarding hypothesis” states that males stay with the female to prevent other males from mating with her. This behavior is advantageous in such situations where mates are scarce and difficult to find. Another explanation is the “male-assistance hypothesis,” where males that remain with a female to help guard and rear their young will have more and healthier offspring. Monogamy is observed in many bird populations where, in addition to the parental care from the female, the male is also a major provider of parental care for the chicks. A third explanation for the evolutionary advantages of monogamy is the “female-enforcement hypothesis.” In this scenario, the female ensures that the male does not have other offspring that might compete with her own, so she actively interferes with the male’s signaling to attract other mates.
Polygynous mating refers to one male mating with multiple females. In these situations, the female must be responsible for most of the parental care as the single male is not capable of providing care to that many offspring. In resourced-based polygyny, males compete for territories with the best resources, and then mate with females that enter the territory, drawn to its resource richness. The female benefits by mating with a dominant, genetically fit male; however, it is at the cost of having no male help in caring for the offspring. An example is seen in the yellow-rumped honeyguide, a bird whose males defend beehives because the females feed on their wax. As the females approach, the male defending the nest will mate with them. Harem mating structures are a type of polygynous system where certain males dominate mating while controlling a territory with resources. Elephant seals, where the alpha male dominates the mating within the group are an example. A third type of polygyny is a lek system. Here there is a communal courting area where several males perform elaborate displays for females, and the females choose their mate from this group. This behavior is observed in several bird species including the sage grouse and the prairie chicken.
In polyandrous mating systems, one female mates with many males. These types of systems are much rarer than monogamous and polygynous mating systems. In pipefishes and seahorses, males receive the eggs from the female, fertilize them, protect them within a pouch, and give birth to the offspring (Figure \(5\)). Therefore, the female is able to provide eggs to several males without the burden of carrying the fertilized eggs.
Simple Learned Behaviors
The majority of the behaviors previously discussed were innate or at least have an innate component (variations on the innate behaviors may be learned). They are inherited and the behaviors do not change in response to signals from the environment. Conversely, learned behaviors, even though they may have instinctive components, allow an organism to adapt to changes in the environment and are modified by previous experiences. Simple learned behaviors include habituation and imprinting—both are important to the maturation process of young animals.
Habituation
Habituation is a simple form of learning in which an animal stops responding to a stimulus after a period of repeated exposure. This is a form of non-associative learning, as the stimulus is not associated with any punishment or reward. Prairie dogs typically sound an alarm call when threatened by a predator, but they become habituated to the sound of human footsteps when no harm is associated with this sound, therefore, they no longer respond to them with an alarm call. In this example, habituation is specific to the sound of human footsteps, as the animals still respond to the sounds of potential predators.
Imprinting
Imprinting is a type of learning that occurs at a particular age or a life stage that is rapid and independent of the species involved. Hatchling ducks recognize the first adult they see, their mother, and make a bond with her. A familiar sight is ducklings walking or swimming after their mothers (Figure \(6\)). This is another type of non-associative learning, but is very important in the maturation process of these animals as it encourages them to stay near their mother so they will be protected, greatly increasing their chances of survival. However, if newborn ducks see a human before they see their mother, they will imprint on the human and follow it in just the same manner as they would follow their real mother.
Link to Learning
The International Crane Foundation has helped raise the world’s population of whooping cranes from 21 individuals to about 600. Imprinting hatchlings has been a key to success: biologists wear full crane costumes so the birds never “see” humans. Watch this video to learn more.
Conditioned Behavior
Conditioned behaviors are types of associative learning, where a stimulus becomes associated with a consequence. During operant conditioning, the behavioral response is modified by its consequences, with regards to its form, strength, or frequency.
Classical Conditioning
In classical conditioning, a response called the conditioned response is associated with a stimulus that it had previously not been associated with, the conditioned stimulus. The response to the original, unconditioned stimulus is called the unconditioned response. The most cited example of classical conditioning is Ivan Pavlov’s experiments with dogs (Figure \(7\)). In Pavlov’s experiments, the unconditioned response was the salivation of dogs in response to the unconditioned stimulus of seeing or smelling their food. The conditioning stimulus that researchers associated with the unconditioned response was the ringing of a bell. During conditioning, every time the animal was given food, the bell was rung. This was repeated during several trials. After some time, the dog learned to associate the ringing of the bell with food and to respond by salivating. After the conditioning period was finished, the dog would respond by salivating when the bell was rung, even when the unconditioned stimulus, the food, was absent. Thus, the ringing of the bell became the conditioned stimulus and the salivation became the conditioned response. Although it is thought by some scientists that the unconditioned and conditioned responses are identical, even Pavlov discovered that the saliva in the conditioned dogs had characteristic differences when compared to the unconditioned dog.
It had been thought by some scientists that this type of conditioning required multiple exposures to the paired stimulus and response, but it is now known that this is not necessary in all cases, and that some conditioning can be learned in a single pairing experiment. Classical conditioning is a major tenet of behaviorism, a branch of psychological philosophy that proposes that all actions, thoughts, and emotions of living things are behaviors that can be treated by behavior modification and changes in the environment.
Operant Conditioning
In operant conditioning, the conditioned behavior is gradually modified by its consequences as the animal responds to the stimulus. A major proponent of such conditioning was psychologist B.F. Skinner, the inventor of the Skinner box. Skinner put rats in his boxes that contained a lever that would dispense food to the rat when depressed. While initially the rat would push the lever a few times by accident, it eventually associated pushing the lever with getting the food. This type of learning is an example of operant conditioning. Operant learning is the basis of most animal training. The conditioned behavior is continually modified by positive or negative reinforcement, often a reward such as food or some type of punishment, respectively. In this way, the animal is conditioned to associate a type of behavior with the punishment or reward, and, over time, can be induced to perform behaviors that they would not have done in the wild, such as the “tricks” dolphins perform at marine amusement park shows (Figure \(8\)).
Cognitive Learning
Classical and operant conditioning are inefficient ways for humans and other intelligent animals to learn. Some primates, including humans, are able to learn by imitating the behavior of others and by taking instructions. The development of complex language by humans has made cognitive learning, the manipulation of information using the mind, the most prominent method of human learning. In fact, that is how students are learning right now by reading this book. As students read, they can make mental images of objects or organisms and imagine changes to them, or behaviors by them, and anticipate the consequences. In addition to visual processing, cognitive learning is also enhanced by remembering past experiences, touching physical objects, hearing sounds, tasting food, and a variety of other sensory-based inputs. Cognitive learning is so powerful that it can be used to understand conditioning in detail. In the reverse scenario, conditioning cannot help someone learn about cognition.
Classic work on cognitive learning was done by Wolfgang Köhler with chimpanzees. He demonstrated that these animals were capable of abstract thought by showing that they could learn how to solve a puzzle. When a banana was hung in their cage too high for them to reach, and several boxes were placed randomly on the floor, some of the chimps were able to stack the boxes one on top of the other, climb on top of them, and get the banana. This implies that they could visualize the result of stacking the boxes even before they had performed the action. This type of learning is much more powerful and versatile than conditioning.
Cognitive learning is not limited to primates, although they are the most efficient in using it. Maze running experiments done with rats by H.C. Blodgett in the 1920s were the first to show cognitive skills in a simple mammal. The motivation for the animals to work their way through the maze was a piece of food at its end. In these studies, the animals in Group I were run in one trial per day and had food available to them each day on completion of the run (Figure \(9\)). Group II rats were not fed in the maze for the first six days and then subsequent runs were done with food for several days after. Group III rats had food available on the third day and every day thereafter. The results were that the control rats, Group I, learned quickly, and figured out how to run the maze in seven days. Group III did not learn much during the three days without food, but rapidly caught up to the control group when given the food reward. Group II learned very slowly for the six days with no reward to motivate them, and they did not begin to catch up to the control group until the day food was given, and then it took two days longer to learn the maze.
It may not be immediately obvious that this type of learning is different than conditioning. Although one might be tempted to believe that the rats simply learned how to find their way through a conditioned series of right and left turns, E.C. Tolman proved a decade later that the rats were making a representation of the maze in their minds, which he called a “cognitive map.” This was an early demonstration of the power of cognitive learning and how these abilities were not just limited to humans.
Sociobiology
Sociobiology is an interdisciplinary science originally popularized by social insect researcher E.O. Wilson in the 1970s. Wilson defined the science as “the extension of population biology and evolutionary theory to social organization.”1 The main thrust of sociobiology is that animal and human behavior, including aggressiveness and other social interactions, can be explained almost solely in terms of genetics and natural selection. This science is controversial; noted scientist such as the late Stephen Jay Gould criticized the approach for ignoring the environmental effects on behavior. This is another example of the “nature versus nurture” debate of the role of genetics versus the role of environment in determining an organism’s characteristics.
Sociobiology also links genes with behaviors and has been associated with “biological determinism,” the belief that all behaviors are hardwired into our genes. No one disputes that certain behaviors can be inherited and that natural selection plays a role retaining them. It is the application of such principles to human behavior that sparks this controversy, which remains active today.
Summary
Behaviors are responses to stimuli. They can either be instinctual/innate behaviors, which are not influenced by the environment, or learned behaviors, which are influenced by environmental changes. Instinctual behaviors include mating systems and methods of communication. Learned behaviors include imprinting and habituation, conditioning, and, most powerfully, cognitive learning. Although the connection between behavior, genetics, and evolution is well established, the explanation of human behavior as entirely genetic is controversial.
Footnotes
1. 1 Edward O. Wilson. On Human Nature (1978; repr., Cambridge: Harvard University Press, 2004), xx.
Glossary
aggressive display
visual display by a species member to discourage other members of the same species or different species
behavior
change in an organism’s activities in response to a stimulus
behavioral biology
study of the biology and evolution of behavior
classical conditioning
association of a specific stimulus and response through conditioning
cognitive learning
knowledge and skills acquired by the manipulation of information in the mind
conditioned behavior
behavior that becomes associated with a specific stimulus through conditioning
courtship display
visual display used to attract a mate
distraction display
visual display used to distract predators away from a nesting site
ethology
biological study of animal behavior
fixed action pattern
series of instinctual behaviors that, once initiated, always goes to completion regardless of changes in the environment
foraging
behaviors species use to find food
habituation
ability of a species to ignore repeated stimuli that have no consequence
imprinting
identification of parents by newborns as the first organism they see after birth
innate behavior
instinctual behavior that is not altered by changes in the environment
intersexual selection
selection of a desirable mate of the opposite sex
intrasexual selection
competition between members of the same sex for a mate
kin selection
sacrificing one’s own life so that one’s genes will be passed on to future generations by relatives
kinesis
undirected movement of an organism in response to a stimulus
learned behavior
behavior that responds to changes in the environment
migration
long-range seasonal movement of animal species
monogamy
mating system whereby one male and one female remain coupled for at least one mating season
operant conditioning
learned behaviors in response to positive and/or negative reinforcement
polyandry
mating system where one female mates with many males
polygyny
mating system where one male mates with many females
reflex action
action in response to direct physical stimulation of a nerve
signal
method of communication between animals including those obtained by the senses of smell, hearing, sight, or touch
taxis
directed movement in response to a stimulus | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/53%3A_Behavioral_Biology/53.01%3A_Natural_History_of_Behavior.txt |
Skills to Develop
• Compare innate and learned behavior
• Discuss how movement and migration behaviors are a result of natural selection
• Discuss the different ways members of a population communicate with each other
• Give examples of how species use energy for mating displays and other courtship behaviors
• Differentiate between various mating systems
• Describe different ways that species learn
Behavior is the change in activity of an organism in response to a stimulus. Behavioral biology is the study of the biological and evolutionary bases for such changes. The idea that behaviors evolved as a result of the pressures of natural selection is not new. Animal behavior has been studied for decades, by biologists in the science of ethology, by psychologists in the science of comparative psychology, and by scientists of many disciplines in the study of neurobiology. Although there is overlap between these disciplines, scientists in these behavioral fields take different approaches. Comparative psychology is an extension of work done in human and behavioral psychology. Ethology is an extension of genetics, evolution, anatomy, physiology, and other biological disciplines. Still, one cannot study behavioral biology without touching on both comparative psychology and ethology.
One goal of behavioral biology is to dissect out the innate behaviors, which have a strong genetic component and are largely independent of environmental influences, from the learned behaviors, which result from environmental conditioning. Innate behavior, or instinct, is important because there is no risk of an incorrect behavior being learned. They are “hard wired” into the system. On the other hand, learned behaviors, although riskier, are flexible, dynamic, and can be altered according to changes in the environment.
Innate Behaviors: Movement and Migration
Innate or instinctual behaviors rely on response to stimuli. The simplest example of this is a reflex action, an involuntary and rapid response to stimulus. To test the “knee-jerk” reflex, a doctor taps the patellar tendon below the kneecap with a rubber hammer. The stimulation of the nerves there leads to the reflex of extending the leg at the knee. This is similar to the reaction of someone who touches a hot stove and instinctually pulls his or her hand away. Even humans, with our great capacity to learn, still exhibit a variety of innate behaviors.
Kinesis and Taxis
Another activity or movement of innate behavior is kinesis, or the undirected movement in response to a stimulus. Orthokinesis is the increased or decreased speed of movement of an organism in response to a stimulus. Woodlice, for example, increase their speed of movement when exposed to high or low temperatures. This movement, although random, increases the probability that the insect spends less time in the unfavorable environment. Another example is klinokinesis, an increase in turning behaviors. It is exhibited by bacteria such as E. coli which, in association with orthokinesis, helps the organisms randomly find a more hospitable environment.
A similar, but more directed version of kinesis is taxis: the directed movement towards or away from a stimulus. This movement can be in response to light (phototaxis), chemical signals (chemotaxis), or gravity (geotaxis) and can be directed toward (positive) or away (negative) from the source of the stimulus. An example of a positive chemotaxis is exhibited by the unicellular protozoan Tetrahymena thermophila. This organism swims using its cilia, at times moving in a straight line, and at other times making turns. The attracting chemotactic agent alters the frequency of turning as the organism moves directly toward the source, following the increasing concentration gradient.
Fixed Action Patterns
A fixed action pattern is a series of movements elicited by a stimulus such that even when the stimulus is removed, the pattern goes on to completion. An example of such a behavior occurs in the three-spined stickleback, a small freshwater fish (Figure \(1\)). Males of this species develop a red belly during breeding season and show instinctual aggressiveness to other males during this time. In laboratory experiments, researchers exposed such fish to objects that in no way resemble a fish in their shape, but which were painted red on their lower halves. The male sticklebacks responded aggressively to the objects just as if they were real male sticklebacks.
Migration
Migration is the long-range seasonal movement of animals. It is an evolved, adapted response to variation in resource availability, and it is a common phenomenon found in all major groups of animals. Birds fly south for the winter to get to warmer climates with sufficient food, and salmon migrate to their spawning grounds. The popular 2005 documentary March of the Penguins followed the 62-mile migration of emperor penguins through Antarctica to bring food back to their breeding site and to their young. Wildebeests (Figure \(2\)) migrate over 1800 miles each year in search of new grasslands.
Although migration is thought of as innate behavior, only some migrating species always migrate (obligate migration). Animals that exhibit facultative migration can choose to migrate or not. Additionally, in some animals, only a portion of the population migrates, whereas the rest does not migrate (incomplete migration). For example, owls that live in the tundra may migrate in years when their food source, small rodents, is relatively scarce, but not migrate during the years when rodents are plentiful.
Foraging
Foraging is the act of searching for and exploiting food resources. Feeding behaviors that maximize energy gain and minimize energy expenditure are called optimal foraging behaviors, and these are favored by natural section. The painted stork, for example, uses its long beak to search the bottom of a freshwater marshland for crabs and other food (Figure \(3\)).
Innate Behaviors: Living in Groups
Not all animals live in groups, but even those that live relatively solitary lives, with the exception of those that can reproduce asexually, must mate. Mating usually involves one animal signaling another so as to communicate the desire to mate. There are several types of energy-intensive behaviors or displays associated with mating, called mating rituals. Other behaviors found in populations that live in groups are described in terms of which animal benefits from the behavior. In selfish behavior, only the animal in question benefits; in altruistic behavior, one animal’s actions benefit another animal; cooperative behavior describes when both animals benefit. All of these behaviors involve some sort of communication between population members.
Communication within a Species
Animals communicate with each other using stimuli known as signals. An example of this is seen in the three-spined stickleback, where the visual signal of a red region in the lower half of a fish signals males to become aggressive and signals females to mate. Other signals are chemical (pheromones), aural (sound), visual (courtship and aggressive displays), or tactile (touch). These types of communication may be instinctual or learned or a combination of both. These are not the same as the communication we associate with language, which has been observed only in humans and perhaps in some species of primates and cetaceans.
A pheromone is a secreted chemical signal used to obtain a response from another individual of the same species. The purpose of pheromones is to elicit a specific behavior from the receiving individual. Pheromones are especially common among social insects, but they are used by many species to attract the opposite sex, to sound alarms, to mark food trails, and to elicit other, more complex behaviors. Even humans are thought to respond to certain pheromones called axillary steroids. These chemicals influence human perception of other people, and in one study were responsible for a group of women synchronizing their menstrual cycles. The role of pheromones in human-to-human communication is still somewhat controversial and continues to be researched.
Songs are an example of an aural signal, one that needs to be heard by the recipient. Perhaps the best known of these are songs of birds, which identify the species and are used to attract mates. Other well-known songs are those of whales, which are of such low frequency that they can travel long distances underwater. Dolphins communicate with each other using a wide variety of vocalizations. Male crickets make chirping sounds using a specialized organ to attract a mate, repel other males, and to announce a successful mating.
Courtship displays are a series of ritualized visual behaviors (signals) designed to attract and convince a member of the opposite sex to mate. These displays are ubiquitous in the animal kingdom. Often these displays involve a series of steps, including an initial display by one member followed by a response from the other. If at any point, the display is performed incorrectly or a proper response is not given, the mating ritual is abandoned and the mating attempt will be unsuccessful. The mating display of the common stork is shown in Figure \(4\).
Aggressive displays are also common in the animal kingdom. An example is when a dog bares its teeth when it wants another dog to back down. Presumably, these displays communicate not only the willingness of the animal to fight, but also its fighting ability. Although these displays do signal aggression on the part of the sender, it is thought that these displays are actually a mechanism to reduce the amount of actual fighting that occurs between members of the same species: they allow individuals to assess the fighting ability of their opponent and thus decide whether it is “worth the fight.” The testing of certain hypotheses using game theory has led to the conclusion that some of these displays may overstate an animal’s actual fighting ability and are used to “bluff” the opponent. This type of interaction, even if “dishonest,” would be favored by natural selection if it is successful more times than not.
Distraction displays are seen in birds and some fish. They are designed to attract a predator away from the nest that contains their young. This is an example of an altruistic behavior: it benefits the young more than the individual performing the display, which is putting itself at risk by doing so.
Many animals, especially primates, communicate with other members in the group through touch. Activities such as grooming, touching the shoulder or root of the tail, embracing, lip contact, and greeting ceremonies have all been observed in the Indian langur, an Old World monkey. Similar behaviors are found in other primates, especially in the great apes.
Link to Learning
The killdeer bird distracts predators from its eggs by faking a broken wing display in this video taken in Boise, Idaho.
Altruistic Behaviors
Behaviors that lower the fitness of the individual but increase the fitness of another individual are termed altruistic. Examples of such behaviors are seen widely across the animal kingdom. Social insects such as worker bees have no ability to reproduce, yet they maintain the queen so she can populate the hive with her offspring. Meerkats keep a sentry standing guard to warn the rest of the colony about intruders, even though the sentry is putting itself at risk. Wolves and wild dogs bring meat to pack members not present during a hunt. Lemurs take care of infants unrelated to them. Although on the surface, these behaviors appear to be altruistic, it may not be so simple.
There has been much discussion over why altruistic behaviors exist. Do these behaviors lead to overall evolutionary advantages for their species? Do they help the altruistic individual pass on its own genes? And what about such activities between unrelated individuals? One explanation for altruistic-type behaviors is found in the genetics of natural selection. In the 1976 book, The Selfish Gene, scientist Richard Dawkins attempted to explain many seemingly altruistic behaviors from the viewpoint of the gene itself. Although a gene obviously cannot be selfish in the human sense, it may appear that way if the sacrifice of an individual benefits related individuals that share genes that are identical by descent (present in relatives because of common lineage). Mammal parents make this sacrifice to take care of their offspring. Emperor penguins migrate miles in harsh conditions to bring food back for their young. Selfish gene theory has been controversial over the years and is still discussed among scientists in related fields.
Even less-related individuals, those with less genetic identity than that shared by parent and offspring, benefit from seemingly altruistic behavior. The activities of social insects such as bees, wasps, ants, and termites are good examples. Sterile workers in these societies take care of the queen because they are closely related to it, and as the queen has offspring, she is passing on genes from the workers indirectly. Thus, it is of fitness benefit for the worker to maintain the queen without having any direct chance of passing on its genes due to its sterility. The lowering of individual fitness to enhance the reproductive fitness of a relative and thus one’s inclusive fitness evolves through kin selection. This phenomenon can explain many superficially altruistic behaviors seen in animals. However, these behaviors may not be truly defined as altruism in these cases because the actor is actually increasing its own fitness either directly (through its own offspring) or indirectly (through the inclusive fitness it gains through relatives that share genes with it).
Unrelated individuals may also act altruistically to each other, and this seems to defy the “selfish gene” explanation. An example of this observed in many monkey species where a monkey will present its back to an unrelated monkey to have that individual pick the parasites from its fur. After a certain amount of time, the roles are reversed and the first monkey now grooms the second monkey. Thus, there is reciprocity in the behavior. Both benefit from the interaction and their fitness is raised more than if neither cooperated nor if one cooperated and the other did not cooperate. This behavior is still not necessarily altruism, as the “giving” behavior of the actor is based on the expectation that it will be the “receiver” of the behavior in the future, termed reciprocal altruism. Reciprocal altruism requires that individuals repeatedly encounter each other, often the result of living in the same social group, and that cheaters (those that never “give back”) are punished.
Evolutionary game theory, a modification of classical game theory in mathematics, has shown that many of these so-called “altruistic behaviors” are not altruistic at all. The definition of “pure” altruism, based on human behavior, is an action that benefits another without any direct benefit to oneself. Most of the behaviors previously described do not seem to satisfy this definition, and game theorists are good at finding “selfish” components in them. Others have argued that the terms “selfish” and “altruistic” should be dropped completely when discussing animal behavior, as they describe human behavior and may not be directly applicable to instinctual animal activity. What is clear, though, is that heritable behaviors that improve the chances of passing on one’s genes or a portion of one’s genes are favored by natural selection and will be retained in future generations as long as those behaviors convey a fitness advantage. These instinctual behaviors may then be applied, in special circumstances, to other species, as long as it doesn’t lower the animal’s fitness.
Finding Sex Partners
Not all animals reproduce sexually, but many that do have the same challenge: they need to find a suitable mate and often have to compete with other individuals to obtain one. Significant energy is spent in the process of locating, attracting, and mating with the sex partner. Two types of selection occur during this process and can lead to traits that are important to reproduction called secondary sexual characteristics: intersexual selection, the choosing of a mate where individuals of one sex choose mates of the other sex, and intrasexual selection, the competition for mates between species members of the same sex. Intersexual selection is often complex because choosing a mate may be based on a variety of visual, aural, tactile, and chemical cues. An example of intersexual selection is when female peacocks choose to mate with the male with the brightest plumage. This type of selection often leads to traits in the chosen sex that do not enhance survival, but are those traits most attractive to the opposite sex (often at the expense of survival). Intrasexual selection involves mating displays and aggressive mating rituals such as rams butting heads—the winner of these battles is the one that is able to mate. Many of these rituals use up considerable energy but result in the selection of the healthiest, strongest, and/or most dominant individuals for mating. Three general mating systems, all involving innate as opposed to learned behaviors, are seen in animal populations: monogamous, polygynous, and polyandrous.
In monogamous systems, one male and one female are paired for at least one breeding season. In some animals, such as the gray wolf, these associations can last much longer, even a lifetime. Several explanations have been proposed for this type of mating system. The “mate-guarding hypothesis” states that males stay with the female to prevent other males from mating with her. This behavior is advantageous in such situations where mates are scarce and difficult to find. Another explanation is the “male-assistance hypothesis,” where males that remain with a female to help guard and rear their young will have more and healthier offspring. Monogamy is observed in many bird populations where, in addition to the parental care from the female, the male is also a major provider of parental care for the chicks. A third explanation for the evolutionary advantages of monogamy is the “female-enforcement hypothesis.” In this scenario, the female ensures that the male does not have other offspring that might compete with her own, so she actively interferes with the male’s signaling to attract other mates.
Polygynous mating refers to one male mating with multiple females. In these situations, the female must be responsible for most of the parental care as the single male is not capable of providing care to that many offspring. In resourced-based polygyny, males compete for territories with the best resources, and then mate with females that enter the territory, drawn to its resource richness. The female benefits by mating with a dominant, genetically fit male; however, it is at the cost of having no male help in caring for the offspring. An example is seen in the yellow-rumped honeyguide, a bird whose males defend beehives because the females feed on their wax. As the females approach, the male defending the nest will mate with them. Harem mating structures are a type of polygynous system where certain males dominate mating while controlling a territory with resources. Elephant seals, where the alpha male dominates the mating within the group are an example. A third type of polygyny is a lek system. Here there is a communal courting area where several males perform elaborate displays for females, and the females choose their mate from this group. This behavior is observed in several bird species including the sage grouse and the prairie chicken.
In polyandrous mating systems, one female mates with many males. These types of systems are much rarer than monogamous and polygynous mating systems. In pipefishes and seahorses, males receive the eggs from the female, fertilize them, protect them within a pouch, and give birth to the offspring (Figure \(5\)). Therefore, the female is able to provide eggs to several males without the burden of carrying the fertilized eggs.
Simple Learned Behaviors
The majority of the behaviors previously discussed were innate or at least have an innate component (variations on the innate behaviors may be learned). They are inherited and the behaviors do not change in response to signals from the environment. Conversely, learned behaviors, even though they may have instinctive components, allow an organism to adapt to changes in the environment and are modified by previous experiences. Simple learned behaviors include habituation and imprinting—both are important to the maturation process of young animals.
Habituation
Habituation is a simple form of learning in which an animal stops responding to a stimulus after a period of repeated exposure. This is a form of non-associative learning, as the stimulus is not associated with any punishment or reward. Prairie dogs typically sound an alarm call when threatened by a predator, but they become habituated to the sound of human footsteps when no harm is associated with this sound, therefore, they no longer respond to them with an alarm call. In this example, habituation is specific to the sound of human footsteps, as the animals still respond to the sounds of potential predators.
Imprinting
Imprinting is a type of learning that occurs at a particular age or a life stage that is rapid and independent of the species involved. Hatchling ducks recognize the first adult they see, their mother, and make a bond with her. A familiar sight is ducklings walking or swimming after their mothers (Figure \(6\)). This is another type of non-associative learning, but is very important in the maturation process of these animals as it encourages them to stay near their mother so they will be protected, greatly increasing their chances of survival. However, if newborn ducks see a human before they see their mother, they will imprint on the human and follow it in just the same manner as they would follow their real mother.
Link to Learning
The International Crane Foundation has helped raise the world’s population of whooping cranes from 21 individuals to about 600. Imprinting hatchlings has been a key to success: biologists wear full crane costumes so the birds never “see” humans. Watch this video to learn more.
Conditioned Behavior
Conditioned behaviors are types of associative learning, where a stimulus becomes associated with a consequence. During operant conditioning, the behavioral response is modified by its consequences, with regards to its form, strength, or frequency.
Classical Conditioning
In classical conditioning, a response called the conditioned response is associated with a stimulus that it had previously not been associated with, the conditioned stimulus. The response to the original, unconditioned stimulus is called the unconditioned response. The most cited example of classical conditioning is Ivan Pavlov’s experiments with dogs (Figure \(7\)). In Pavlov’s experiments, the unconditioned response was the salivation of dogs in response to the unconditioned stimulus of seeing or smelling their food. The conditioning stimulus that researchers associated with the unconditioned response was the ringing of a bell. During conditioning, every time the animal was given food, the bell was rung. This was repeated during several trials. After some time, the dog learned to associate the ringing of the bell with food and to respond by salivating. After the conditioning period was finished, the dog would respond by salivating when the bell was rung, even when the unconditioned stimulus, the food, was absent. Thus, the ringing of the bell became the conditioned stimulus and the salivation became the conditioned response. Although it is thought by some scientists that the unconditioned and conditioned responses are identical, even Pavlov discovered that the saliva in the conditioned dogs had characteristic differences when compared to the unconditioned dog.
It had been thought by some scientists that this type of conditioning required multiple exposures to the paired stimulus and response, but it is now known that this is not necessary in all cases, and that some conditioning can be learned in a single pairing experiment. Classical conditioning is a major tenet of behaviorism, a branch of psychological philosophy that proposes that all actions, thoughts, and emotions of living things are behaviors that can be treated by behavior modification and changes in the environment.
Operant Conditioning
In operant conditioning, the conditioned behavior is gradually modified by its consequences as the animal responds to the stimulus. A major proponent of such conditioning was psychologist B.F. Skinner, the inventor of the Skinner box. Skinner put rats in his boxes that contained a lever that would dispense food to the rat when depressed. While initially the rat would push the lever a few times by accident, it eventually associated pushing the lever with getting the food. This type of learning is an example of operant conditioning. Operant learning is the basis of most animal training. The conditioned behavior is continually modified by positive or negative reinforcement, often a reward such as food or some type of punishment, respectively. In this way, the animal is conditioned to associate a type of behavior with the punishment or reward, and, over time, can be induced to perform behaviors that they would not have done in the wild, such as the “tricks” dolphins perform at marine amusement park shows (Figure \(8\)).
Cognitive Learning
Classical and operant conditioning are inefficient ways for humans and other intelligent animals to learn. Some primates, including humans, are able to learn by imitating the behavior of others and by taking instructions. The development of complex language by humans has made cognitive learning, the manipulation of information using the mind, the most prominent method of human learning. In fact, that is how students are learning right now by reading this book. As students read, they can make mental images of objects or organisms and imagine changes to them, or behaviors by them, and anticipate the consequences. In addition to visual processing, cognitive learning is also enhanced by remembering past experiences, touching physical objects, hearing sounds, tasting food, and a variety of other sensory-based inputs. Cognitive learning is so powerful that it can be used to understand conditioning in detail. In the reverse scenario, conditioning cannot help someone learn about cognition.
Classic work on cognitive learning was done by Wolfgang Köhler with chimpanzees. He demonstrated that these animals were capable of abstract thought by showing that they could learn how to solve a puzzle. When a banana was hung in their cage too high for them to reach, and several boxes were placed randomly on the floor, some of the chimps were able to stack the boxes one on top of the other, climb on top of them, and get the banana. This implies that they could visualize the result of stacking the boxes even before they had performed the action. This type of learning is much more powerful and versatile than conditioning.
Cognitive learning is not limited to primates, although they are the most efficient in using it. Maze running experiments done with rats by H.C. Blodgett in the 1920s were the first to show cognitive skills in a simple mammal. The motivation for the animals to work their way through the maze was a piece of food at its end. In these studies, the animals in Group I were run in one trial per day and had food available to them each day on completion of the run (Figure \(9\)). Group II rats were not fed in the maze for the first six days and then subsequent runs were done with food for several days after. Group III rats had food available on the third day and every day thereafter. The results were that the control rats, Group I, learned quickly, and figured out how to run the maze in seven days. Group III did not learn much during the three days without food, but rapidly caught up to the control group when given the food reward. Group II learned very slowly for the six days with no reward to motivate them, and they did not begin to catch up to the control group until the day food was given, and then it took two days longer to learn the maze.
It may not be immediately obvious that this type of learning is different than conditioning. Although one might be tempted to believe that the rats simply learned how to find their way through a conditioned series of right and left turns, E.C. Tolman proved a decade later that the rats were making a representation of the maze in their minds, which he called a “cognitive map.” This was an early demonstration of the power of cognitive learning and how these abilities were not just limited to humans.
Sociobiology
Sociobiology is an interdisciplinary science originally popularized by social insect researcher E.O. Wilson in the 1970s. Wilson defined the science as “the extension of population biology and evolutionary theory to social organization.”1 The main thrust of sociobiology is that animal and human behavior, including aggressiveness and other social interactions, can be explained almost solely in terms of genetics and natural selection. This science is controversial; noted scientist such as the late Stephen Jay Gould criticized the approach for ignoring the environmental effects on behavior. This is another example of the “nature versus nurture” debate of the role of genetics versus the role of environment in determining an organism’s characteristics.
Sociobiology also links genes with behaviors and has been associated with “biological determinism,” the belief that all behaviors are hardwired into our genes. No one disputes that certain behaviors can be inherited and that natural selection plays a role retaining them. It is the application of such principles to human behavior that sparks this controversy, which remains active today.
Summary
Behaviors are responses to stimuli. They can either be instinctual/innate behaviors, which are not influenced by the environment, or learned behaviors, which are influenced by environmental changes. Instinctual behaviors include mating systems and methods of communication. Learned behaviors include imprinting and habituation, conditioning, and, most powerfully, cognitive learning. Although the connection between behavior, genetics, and evolution is well established, the explanation of human behavior as entirely genetic is controversial.
Footnotes
1. 1 Edward O. Wilson. On Human Nature (1978; repr., Cambridge: Harvard University Press, 2004), xx.
Glossary
aggressive display
visual display by a species member to discourage other members of the same species or different species
behavior
change in an organism’s activities in response to a stimulus
behavioral biology
study of the biology and evolution of behavior
classical conditioning
association of a specific stimulus and response through conditioning
cognitive learning
knowledge and skills acquired by the manipulation of information in the mind
conditioned behavior
behavior that becomes associated with a specific stimulus through conditioning
courtship display
visual display used to attract a mate
distraction display
visual display used to distract predators away from a nesting site
ethology
biological study of animal behavior
fixed action pattern
series of instinctual behaviors that, once initiated, always goes to completion regardless of changes in the environment
foraging
behaviors species use to find food
habituation
ability of a species to ignore repeated stimuli that have no consequence
imprinting
identification of parents by newborns as the first organism they see after birth
innate behavior
instinctual behavior that is not altered by changes in the environment
intersexual selection
selection of a desirable mate of the opposite sex
intrasexual selection
competition between members of the same sex for a mate
kin selection
sacrificing one’s own life so that one’s genes will be passed on to future generations by relatives
kinesis
undirected movement of an organism in response to a stimulus
learned behavior
behavior that responds to changes in the environment
migration
long-range seasonal movement of animal species
monogamy
mating system whereby one male and one female remain coupled for at least one mating season
operant conditioning
learned behaviors in response to positive and/or negative reinforcement
polyandry
mating system where one female mates with many males
polygyny
mating system where one male mates with many females
reflex action
action in response to direct physical stimulation of a nerve
signal
method of communication between animals including those obtained by the senses of smell, hearing, sight, or touch
taxis
directed movement in response to a stimulus
53.03: Behavioral Genetics
Behavior is action that alters the relationship between an organism and its environment. Behavior may occur as a result of
• an external stimulus (e.g., sight of a predator)
• internal stimulus (e.g., hunger)
• or, more often, a mixture of the two (e.g., mating behavior)
It is often useful to distinguish between
• innate behavior = behavior determined by the "hard-wiring" of the nervous system. It is usually inflexible, a given stimulus triggering a given response. A salamander raised away from water until long after its siblings begin swimming successfully will swim every bit as well as they the very first time it is placed in the water. Clearly this rather elaborate response is "built in" in the species and not something that must be acquired by practice.
• learned behavior = behavior that is more or less permanently altered as a result of the experience of the individual organism (e.g., learning to play baseball well).
• However, careful analysis often reveals that any particular behavior is a combination of innate and learned components.
Examples of innate behavior:
• taxes
• reflexes
• instincts
Taxes
Some organisms respond to a stimulus by automatically moving directly toward or away from or at some defined angle to it. These responses are called taxes. They are similar to tropisms in plants except that actual locomotion of the entire organism is involved. Link to a detailed discussion.
Reflexes
The Withdrawal Reflex
When you touch a hot object, you quickly pull you hand away using the withdrawal reflex.
These are the steps:
• The stimulus is detected by receptors in the skin.
• These initiate nerve impulses in sensory neurons leading from the receptors to the spinal cord.
• The impulses travel into the spinal cord where the sensory nerve terminals synapse with interneurons.
• Some of these synapse with motor neurons that travel out from the spinal cord entering mixed nerves that lead to the flexors that withdraw your hand.
• Others synapse with inhibitory interneurons that suppress any motor output to extensors whose contraction would interfere with the withdrawal reflex.
The Stretch Reflex
The stretch reflex is examined (with a diagram) on a separate page. Link to it.
Instincts
Instincts are complex behavior patterns which, like reflexes, are inborn, rather inflexible, and valuable at adapting the animal to its environment. They differ from reflexes in their complexity. The entire body participates in instinctive behavior, and an elaborate series of actions may be involved.
The scratching behavior of a dog and a European bullfinch, shown here, is part of their genetic heritage. The widespread behavior of scratching with a hind limb crossed over a forelimb in common to most birds, reptiles, and mammals. (Picture courtesy of Rudolf Freund and Scientific American, 1958.) So instincts are inherited just as the structure of tissues and organs is. Another example.
• The African peach-faced lovebird carries nesting materials to the nesting site by tucking them in its feathers.
• Its close relative, the Fischer's lovebird, uses its beak to transport nesting materials.
• The two species can hybridize. When they do so, the offspring succeed only in carrying nesting material in their beaks. Nevertheless, they invariably go through the motions of trying to tuck the materials in their feathers first.
Foraging Behavior
Foraging for food is a crucial behavior for animals. Like all behavior, it requires the interaction of many components. Nonetheless, it turns out that in some animals, at least, foraging behavior can be altered by a single gene.
Drosophila melanogaster
The discovery of the genetic control of foraging in Drosophila began with the observations of Marla Sokolowski when she was an undergraduate biology student at the University of Toronto.
She noticed that Drosophila larvae, feeding in her culture vessels, displayed one of two distinct feeding patterns:
• "rovers" who moved rapidly over the surface of the culture medium
• "sitters who fed at a much more leisurely pace
She went on to find that this "bimodal" pattern of behavior
• continued when the larvae became adults
• was present in populations of wild fruit flies, not just in her laboratory colonies
After further years of research, she has shown that the behavior is under the control of a single gene, named for ("foraging"). Two alleles are present, at almost equal frequencies, that is, for is polymorphic.
• forR, which is dominant
• fors, the recessive
• About 70% of natural populations are rovers being either homozygous for forR or heterozygous (forR/fors).
• Sitters are homozygous for fors
Both alleles encode a PKG, a protein kinase (an enzyme that attaches phosphate groups to target proteins) that is activated by the "second messenger" cyclic GMP (cGMP). The enzyme encoded by the forR allele is more active than that encoded by fors. She and her colleagues have succeeded in inserting forR DNA into sitters who promptly become rovers.
Why this polymorphism? Why should alleles for two such different behaviors be maintained at such high frequency in the population?
One possible answer: it permits the population to thrive under varying food conditions:
• sitters are favored when food is abundant
• rovers are favored when competition for food is strong, such as in crowded cultures
Honeybees
Honeybees have their version of the for gene, called Amfor ("Apis mellifera for"). It, too, encodes a cGMP-dependent protein kinase (PKG). When worker bees first hatch, they remain in the hive tending to various housekeeping chores, such as feeding the larvae. But when they are 2–3 weeks old, they leave the hive and begin foraging for nectar and pollen. This change in behavior coincides with the increased expression of Amfor. When newly-hatched workers are treated with cGMP, the amount of PKG in their brains goes up and they quickly begin foraging instead of doing housekeeping.
Interaction of Internal and External Stimuli
Instinctive behavior often depends on conditions in the internal environment. In many vertebrates courtship and mating behavior will not occur unless sex hormones (estrogens in females, androgens in males) are present in the blood. The target organ is a small region of the hypothalamus. When stimulated by sex hormones in its blood supply, the hypothalamus initiates the activities leading to mating. The level of sex hormones is, in turn, regulated by the activity of the anterior lobe of the pituitary gland.
The above figure outlines the interactions of external and internal stimuli that lead an animal, such as a rabbit, to see a sexual partner and mate with it.
Releasers of Instinctive Behavior
Once the body is prepared for certain types of instinctive behavior, an external stimulus may be needed to initiate the response. N. Tinbergen (who shared the 1973 Nobel Prize with Konrad Lorenz and Karl von Frisch) showed that the stimulus need not necessarily be appropriate to be effective.
• During the breeding season, the female three-spined stickleback normally follows the red-bellied male (a in the figure) to the nest that he has prepared.
• He guides her into the nest (b) and then
• prods the base of her tail (c).
• She then lays eggs in the nest.
• After doing so, the male drives her from the nest, enters it himself, and fertilizes the eggs (d).
• Although this is the normal pattern, the female will follow almost any small red object to the nest, and once within the nest, neither the male nor any other red object need be present.
• Any object touching her near the base of her tail will cause her to release her eggs.
It is as though she were primed internally for each item of behavior and needed only one specific signal to release the behavior pattern. For this reason, signals that trigger instinctive acts are called releasers. Once a particular response is released, it usually runs to completion even though the stimulus has been removed. One or two prods at the base of her tail will release the entire sequence of muscular actions involved in liberating her eggs.
Chemical signals (e.g., pheromones) serve as important releasers for the social insects: ants, bees, and termites. Many of these animals emit several different pheromones which elicit, for example, alarm behavior, mating behavior, and foraging behavior in other members of their species.
The mammary glands of domestic rabbit mothers emit a pheromone that releases immediate nursing behavior by their babies (pups). A good thing, too, as mothers devote only 5–7 minutes a day to feeding their pups so they had better be quick about it.
The studies of Tinbergen and others have shown that animals can often be induced to respond to inappropriate releasers. For example, a male robin defending its territory will repeatedly attack a simple clump of red feathers instead of a stuffed robin that lacks the red breast of the males.
Although such behavior seems inappropriate to our eyes, it reveals a crucial feature of all animal behavior: animals respond selectively to certain aspects of the total sensory input they receive. Animals spend their lives bombarded by myriad sights, sounds, odors, etc. But their nervous system filters this mass of sensory data, and they respond only to those aspects that the evolutionary history of the species has proved to be significant for survival. | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/53%3A_Behavioral_Biology/53.03%3A_Behavioral_Genetics/53.3.01%3A_Innate_Behavior.txt |
Habituation
Habituation is a reduction in a previously-displayed response when no reward or punishment follows. If you make an unusual sound in the presence of the family dog, it will respond usually by turning its head toward the sound. If the stimulus is given repeatedly and nothing either pleasant or unpleasant happens to the dog, it will soon cease to respond. This lack of response is not a result of fatigue or sensory adaptation and is long-lasting; when fully habituated, the dog will not respond to the stimulus even though weeks or months have elapsed since it was last presented.
Sensitization
Sensitization is an increase in the response to an innocuous stimulus when that stimulus occurs after a punishing stimulus.
Sensitization in sea slugs
When the siphon of the sea slug Aplysia is gently touched, the animal withdraws its gill for a brief period. However, if preceded by an electrical shock to its tail, the same gentle touch to the siphon will elicit a longer period of withdrawal. The sensitization response to a single shock (blue bar) dies out after about an hour, and returns to baseline after a day (yellow). So it is an example of short-term memory.
However, if the animal is sensitized with multiple shocks given over several days, its subsequent response to a gentle touch on the siphon is much larger and is retained longer (tan and gray bars). This is an example of long-term memory and requires protein synthesis. (These findings are the work of Eric R. Kandel, who was awarded a Nobel Prize in 2000.)
Imprinting
If newly-hatched geese are exposed to a moving object of reasonable size and emitting reasonable sounds, they will begin to follow it just as they would normally follow their mother. This is called imprinting. The time of exposure is quite critical. A few days after hatching, imprinting no longer occurs. Prior to this time, though, the results can be quite remarkable. A gosling imprinted to a moving box or clucking person will try to follow this object for the rest of its life. In fact, when the gosling reaches sexual maturity, it will make the imprinted object - rather than a member of its own species - the goal of its sexual drive.
Much of our knowledge of imprinting was learned from the research of Konrad Lorenz, shown here with some of his imprinted goslings. Lorenz shared a Nobel Prize in 1973 for his discoveries. (Photo by Tom McAvoy; courtesy of LIFE Magazine, ©1955, Time, Inc.)
Male mice become imprinted with the odor of littermates during the first three weeks of life. When they reach sexual maturity, they avoid mating with close relatives. The odor is controlled by the major histocompatibility complex (MHC).
The Conditioned Response (CR)
The conditioned response is probably the simplest form of learned behavior. It is a response that as a result of experience comes to be caused by a stimulus different from the one that originally triggered it. The Russian physiologist Ivan Pavlov found that placing meat powder in a dog's mouth would cause it to salivate.
The meat powder, an unconditioned stimulus (US), triggers a simple inborn reflex involving taste receptors, sensory neurons, networks of interneurons in the brain, and autonomic motor neurons running to the salivary glands thus producing an unconditioned response (UR).
Pavlov found that if he rang a bell every time he put the meat powder in the dog's mouth, the dog eventually salivated upon hearing the bell alone. This is the conditioned response (CR). The dog has learned to respond to a substitute stimulus, the conditioned stimulus (CS).
We assume that the physiological basis of the conditioned response is the transfer, by appropriate neurons, of nervous activity in the auditory areas of the brain to the motor neurons controlling salivation. This involves the development and/or strengthening of neural circuits, which - we may also assume - is characteristic of all forms of learning.
The conditioned response has proved to be an excellent tool for determining the sensory capabilities of other animals. For example, honeybees can be conditioned to seek food on a piece of blue cardboard. By offering other colors to a blue-conditioned bee, Karl von Frisch (who shared the 1973 Nobel Prize with Lorenz) found that honeybees can discriminate between yellow-green, blue-green, blue-violet, and ultraviolet.
Instrumental Conditioning
Pavlov's dogs were restrained and the response being conditioned (salivation) was innate. But the principles of conditioning can also be used to train animals to perform tasks that are not innate. In these cases, the animal is placed in a setting where it can move about and engage in different activities. The experimenter chooses to reward only one, e.g., turning to the left. By first rewarding (e.g., with a pellet of food) even the slightest movement to the left and then only more complete turns, a skilled experimenter can in about 2 minutes train a naive pigeon to make a complete turn. A little more work and the pigeon will pace out a figure eight.
In the example shown here, the pigeon - presented with two spots of light - pecks at the brighter and reaches down to pick up the grain of food that is its reward.
Such training is known as instrumental conditioning or operant conditioning. The latter term was coined by B. F. Skinner, whose skill with the technique enabled him to train pigeons to play ping-pong and even a toy piano! It is also called trial-and-error learning because the animal is free to try various responses before finding the one that is rewarded.
Maze problems are a form of instrumental conditioning in which the animal is faced with a sequence of alternatives.
In this photo (Courtesy of B. Rensch), Julia, a chimpanzee, uses a magnet to move an iron ring through a maze. Julia is able to solve mazes like this on her first attempt most (86%) of the time and sometimes faster than biology students can!
Concepts
Although most animals solve mazes and other problems by trial and error, Julia (and biology students) usually make only one or two random attempts at solving a problem and then, all of a sudden, "get it". They have made an abstract generalization about the specific problem; that is, have formed a concept. Oddity problems are an example. This young rhesus monkey has learned that food will be found - not under any particular object - but under whichever object is different from the others.
In monkeys (and probably humans as well), concept formation depends on activity in the prefrontal cortex of the brain. Recent research suggests that honeybees can also solve simple oddity problems! | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/53%3A_Behavioral_Biology/53.04%3A_Learning.txt |
The domestic honeybee, Apis mellifera, is a colonial insect living in hives containing one queen — a fertile female, a few drones (males), and thousands of workers (infertile females). The workers are responsible for keeping the hive clean, building the wax combs of the hive, tending the young and, when they get older (and when their for gene gets turned on), and foraging for food - nectar and pollen.
ome 5–25% of the workers in the hive are scouts. Their job is to search for new sources of food for the other workers, the foragers, to harvest. While both scouts and foragers look alike, recent research suggests that they represent stable subpopulations with distinctive patterns of gene expression in their brains. (See Liang, Z. S., et al., in the 9 March 2012 issue of Science.)
When food is discovered by scouts, they return to the hive. Shortly after their return, many foragers leave the hive and fly directly to the food. The remarkable thing about this is that the foragers do not follow the scouts back. The scouts may remain in the hive for hours and those that leave continue to hunt for new sources of food even though the foragers are continuing to bring back ample supplies of food from the sites the scouts discovered earlier. So the scout bees have communicated to the foragers the necessary information for them to find the food on their own.
It turns out that the scouts can convey to the foragers information about
• the odor of the food
• its direction from the hive
• its distance from the hive
Distance
When food is within 50–75 meters of the hive, the scouts dance the "round dance" on the surface of the comb (left). But when the food is farther than 75 meters from the hive, the scouts dance the "waggle dance" (right). The waggle dance has two components: (1) a straight run - the direction of which conveys information about the direction of the food and (2) the speed at which the dance is repeated which indicates how far away the food is.
The graph shows the relationship between the speed of the dance and the distance to the food. It is based on data collected by the German ethologist Karl von Frisch. It was he who discovered much of what we know today about honeybee communication (and was honored with a Nobel Prize in 1973).
How do the bees calculate distance? von Frisch thought they measured the distance by assessing the amount of energy it took to get there. But the mechanism turns out to be quite different. The bees measure distance by the motion of images received by their eyes as they fly.
Flicker Effect
Honeybees, like all insects, have compound eyes. These give little information about depth but are very sensitive to "flicker effect". It has long been known to bee keepers that honeybees respond better to flowers
• moving in the breeze
• with complex petals
The importance of flicker effect can be demonstrated by training honeybees to visit food placed on cards with patterns. For example, the bees can distinguish any figure in the top row from any figure in the bottom row more easily than they can distinguish between any of the figures in either row.
Evidence of distance measuring
Working at the University of Notre Dame (Indiana), Esch and Burns found that when the hive and food were placed on top of tall (50 m) buildings, the speed of the waggle dance indicated a distance only half of the distance indicated by bees travelling the same distance (230 m) at ground level. Features of the scenery pass the retina faster when near than when far (compare the changes in your view from an airliner at cruising altitude and as it approaches the runway).
Tests of Searching Behavior
Working at the Australian National University, Srinivasan and his colleagues built tunnels decorating the interior walls with patterns to create flicker. In these three experiments, the bees were first fed in the middle of the tunnel of standard diameter. Then the food was removed.
1. Using the same tunnel, foragers came to same spot in the tunnel.
2. Using a narrower tunnel, foragers began searching for food near its entrance. Forced to fly closer to the vertical stripes, the images moved by faster.
3. Using a tunnel with a larger diameter, foragers searched for food at its far end. Flying farther from the stripes, the images moved by more slowly.
Tests of Dancing Behavior
All tunnels were 6 meters long and located 35 meters from the hive — well within the distance (50–75 m) that normally elicits the round dance.
1. Bees were fed at the entrance. Back at the hive, they danced the round dance as you would expect.
2. Bees fed at end of tunnel. Back at the hive, they danced the waggle dance. Even though the total distance from the hive (41 m) was still well within the "round" range, the complexity of the scenery passed in the last 6 m, elicited the waggle dance.
3. Bees fed at end of tunnel decorated with horizontal stripes. These created no flicker as the bees flew, and on their return to the hive they danced the round dance.
You can read about this second set of experiments in Srinivasan, et. al., Science, 4 February 2000.
Recruits respond to the misinformation given by returning scouts
More recently, the Esch and Srinivasan groups have teamed up to show that naive foragers are fooled by the misinformation that the returning scouts give them. When scouts were fed only 11 m from the hive but at the far end of a striped tunnel, they danced the waggle dance back at the hive as though the food had been 70 m away. Most of the workers they recruited flew out (bypassing the tunnel) 70 m looking for food. You can read about these experiments in Ungless, et. al., Nature, 31 May 2001.
Direction
By itself, the knowledge that food is 6 kilometers (3.7 miles) away is not very useful. But von Frisch also noted that the direction of the straight portion of the waggle dance varied with the direction of the food source from the hive and the time of day.
• At any one time, the direction changes with the location of the food.
• With a fixed source of food, the direction changes by the same angle as the sun during its passage through the sky.
But
• The sun is not visible within the hive.
• The scouts dance on the vertical surface of the combs.
How, then, do they translate flight angles in the darkened hive?
The picture shows the relationship between the angle of the dance on the vertical comb and the bearing of the sun with respect to the location of food. When the food and sun are in the same direction, the straight portion of the waggle dance is directed upward. When the food is at some angle to the right (blue) or left (red) of the sun, the bee orients the straight portion of her dance at the same angle to the right or left of the vertical.
Using radar to track individual bees recruited by the waggle dance, Riley, J. R., et al., (Nature, 12 May 2005) have shown that the recruits do fly in the indicated direction. They even adjust their flight path to compensate for being blown off course by the wind. However, their course is seldom so precise that they can find the food without the aid of vision and/or smell as they neared it.
Other features
Time Sense
When scouts remain in the hive for a long period, they shift the direction of the straight portion of the waggle dance as the day wears on (and the direction of the sun shifts). But they cannot see the sun in the darkened hive. Evidently, they are "aware" of the passing time and make the necessary corrections.
The time sense of honeybees has long been known to people who have sweet snacks in their garden at a set time every day. Within minutes of the regular time, foraging bees arrive for their share of the jam. The speed of the bee's clock seems to be related to its metabolic rate. If normally punctual bees are chilled (to lower their metabolic rate) or exposed to an anesthetizing concentration of carbon dioxide they arrive late to the picnic table (graph above).
Polarized light
von Frisch also discovered that scouts (and foragers) don't actually have to see the sun to navigate. As long as they can see a small patch of clear blue sky, they get along fine. This is because sky light is partially polarized, and the plane of polarization in any part of the sky is determined by the location of the sun. Try it by rotating a pair of polaroid® sun glasses!
Swarming
Before a new queen emerges, the old queen leaves the hive, taking many of the workers with her. The swarm usually settles somewhere, e.g., on a tree branch, while scouts go searching for a new home. Each scout that finds a promising site, returns to the swarm and dances on it just as though she had found food. Eventually, the swarm departs for the location promoted most vigorously. Once the new hive is established, many of the scouts that found the site will become food scouts.
Grooming
Workers have another type of dance — rapidly vibrating from side to side — that tells other workers that she needs help removing dust, pollen, etc. from hard-to-reach places on her body. | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/53%3A_Behavioral_Biology/53.08%3A_Animal_Communication/53.8.01%3A_Honeybee_Navigation.txt |
Pheromones are chemicals released by an organism into its environment enabling it to communicate with other members of its own species.
Pheromones in Insects
Alarm Pheromone: When an ant is disturbed, it releases a pheromone that can be detected by other ants several centimeters away. They are attracted by low concentrations of the pheromone and begin to move toward the region of increasing concentration. As they get nearer to their disturbed nestmate, their response changes to one of alarm. The higher concentration causes them to run about as they work to remedy the disturbance. Unless additional amounts of the alarm pheromone are released, it soon dissipates. This ensures that once the emergency is over, the ants return quietly to their former occupations. Honeybees also have an alarm pheromone (which is a good thing not to elicit around a colony of "Africanized" bees).
Trail Pheromone: Certain ants, as they return to the nest with food, lay down a trail pheromone. This trail attracts and guides other ants to the food. It is continually renewed as long as the food holds out. When the supply begins to dwindle, trailmaking ceases. The trail pheromone evaporates quickly so other ants stop coming to the site and are not confused by old trails when food is found elsewhere. And at least in one species of ant, trails that no longer lead to food are also marked with a repellant pheromone.
A stick treated with the trail pheromone of an ant (left) can be used to make an artificial trail which is followed closely by other ants emerging from their nest (right). The trail will not be maintained by other ants unless food is placed at its end. (Photos courtesy of Sol Mednick and Scientific American).
Queen Pheromone: Honeybee queens spend their lives literally surrounded by a retinue of worker bees. The queen emits a pheromone that is a complex mixture of unsaturated hydrocarbons, fatty acids, and other organic molecules. Among its effects:
• inducing the workers to feed and groom her
• inhibiting the workers from building queen cells and rearing new queens
• inhibiting ovary development in the workers
Sex Attractants
Hundreds of pheromones are known with which one sex (usually the female) of an insect species attracts its mates. Many of these sex attractants or their close chemical relatives are available commercially. They have proved useful weapons against insect pests in two ways:
• Male Confusion: Distributing a sex attractant throughout an area masks the insect's own attractant and thus may prevent the sexes getting together. This "communication disruption" has been used successfully against a wide variety of important pests. For example, the sex attractant of the cotton boll weevil has reduced the need for conventional chemical insecticides by more than half in some cotton-growing areas.
• Insect Monitoring: Insect sex attractants are also valuable in monitoring pest populations. By baiting traps with the appropriate pheromone, a build-up of the pest population can be spotted early. Even if a conventional insecticide is the weapon chosen, its early use reduces
• the amount needed
• damage to the crop
• cost to the grower
• possible damage to the environment.
Early detection of pest build-up is a key ingredient in the system known as integrated pest management (IPM).
The photo (courtesy of USDA) shows the feathery antennae of a male gypsy moth. These detect the pheromone released by the females (who do not fly). In some insects, a single molecule of sex attractant is enough to elicit a response.
Sexual Deception
• By an animal: Many species of spiders prey exclusively on moths of certain species and only on the males. Studies of one species of spider, Mastophora cornigera, show that it releases a mixture of volatile compounds that mimic the sex pheromone of the moth species it preys upon. Male moths flying upwind in search of a female end up eaten instead!
• By a plant: A number of species of orchids - each pollinated by the males of a single species of insect (wasps or bees) — emit the same pheromone that is the sex attractant by which females of the insect species attract the males for mating.
Pheromones in Mammals
Releaser Pheromones: Many mammals (e.g., dogs and cats) deposit chemicals in and/or around their "territory". As these vaporize, they signal to other members of the species of the presence of the occupant of the territory. Domestic rabbit mothers release a mammary pheromone that triggers immediate nursing behavior by their babies (pups). A good thing, too, as mothers devote only 5–7 minutes a day to feeding their pups so they had better be quick about it.
Many animals, including mammals, signal with alarm pheromones. Although neither the source nor the chemical nature of alarm pheromones are known in any mammal, stressed animals release something that triggers quick behavior (e.g., flight or fight) in other members of their species. The pheromone is detected in a special cluster of cells located at the very tip of the nose and thus in a position to detected airborne molecules even before the vomeronasal organ (see below) or nasal epithelium can. The detectors on these cells are primary cilia.
Primer Pheromones: Rats and mice give off pheromones that elicit mating behavior. However, the response is not immediate as it is in the releaser pheromones of mother rabbits and insects. Instead, detection of the pheromone primes the endocrine system of the recipient to make the changes, e.g., ovulation, needed for successful mating. Primer pheromones are detected by the olfactory epithelium with which normal odors are detected and also in most mammals (but not humans) by the vomeronasal organ (VNO). The VNO is a patch of receptor tissue in the nasal cavity distinct from the olfactory epithelium. The receptors are G-protein-coupled transmembrane proteins similar to those that mediate olfaction, but encoded by entirely different genes. The neurons leading from the VNO take a separate path into and through the brain.
Human pheromones: It has long been noticed that women living close together (e.g., college roommates) develop synchronous menstrual cycles. This is thought to be because they release two (as yet uncharacterized) primer pheromones: (1) one prior to ovulation that tends to speed up the onset of ovulation in others and (2) one after ovulation that tends to delay the onset of ovulation in other women. Both pheromones are released from the armpits. The pheromones are not detected consciously as odors, but presumably trigger the hormonal changes that mediate the menstrual cycle. | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/53%3A_Behavioral_Biology/53.08%3A_Animal_Communication/53.8.02%3A_Pheromones.txt |
Imagine sailing down a river in a small motorboat on a weekend afternoon; the water is smooth and you are enjoying the warm sunshine and cool breeze when suddenly you are hit in the head by a 20-pound silver carp. This is a risk now on many rivers and canal systems in Illinois and Missouri because of the presence of Asian carp.
This fish—actually a group of species including the silver, black, grass, and big head carp—has been farmed and eaten in China for over 1000 years. It is one of the most important aquaculture food resources worldwide. In the United States, however, Asian carp is considered a dangerous invasive species that disrupts community structure and composition to the point of threatening native species.
54.2.01: Prelude to Population and Community Ecolo
Imagine sailing down a river in a small motorboat on a weekend afternoon; the water is smooth and you are enjoying the warm sunshine and cool breeze when suddenly you are hit in the head by a 20-pound silver carp. This is a risk now on many rivers and canal systems in Illinois and Missouri because of the presence of Asian carp.
This fish—actually a group of species including the silver, black, grass, and big head carp—has been farmed and eaten in China for over 1000 years. It is one of the most important aquaculture food resources worldwide. In the United States, however, Asian carp is considered a dangerous invasive species that disrupts community structure and composition to the point of threatening native species.
54.03: Population Demography and Dynamics
• 54.1: The Environmental Challenges
• 54.2: Populations, Groups of Single Species in One Place
• 54.3: Population Demography and Dynamics
Populations are dynamic entities. Populations consist all of the species living within a specific area, and populations fluctuate based on a number of factors: seasonal and yearly changes in the environment, natural disasters such as forest fires and volcanic eruptions, and competition for resources between and within species. The statistical study of population dynamics, demography, uses a series of mathematical tools to investigate how populations respond to changes in their environments.
• 54.4: Life History and the Cost of Reproduction
A species’ life history describes the series of events over its lifetime, such as how resources are allocated for growth, maintenance, and reproduction. Life history traits affect the life table of an organism. A species’ life history is genetically determined and shaped by the environment and natural selection.
• 54.5: Environmental Limits to Population Growth
Although life histories describe the way many characteristics of a population (such as their age structure) change over time in a general way, population ecologists make use of a variety of methods to model population dynamics mathematically. These more precise models can then be used to accurately describe changes occurring in a population and better predict future changes.
• 54.6: Factors that Regulate Populations
The logistic model of population growth, while valid in many natural populations and a useful model, is a simplification of real-world population dynamics. Implicit in the model is that the carrying capacity of the environment does not change, which is not the case. The carrying capacity varies annually: for example, some summers are hot and dry whereas others are cold and wet. In many areas, the carrying capacity during the winter is much lower than it is during the summer.
• 54.7: Human Population Growth
Although humans have increased the carrying capacity of their environment, the technologies used to achieve this transformation have caused unprecedented changes to Earth’s environment, altering ecosystems to the point where some may be in danger of collapse. The depletion of the ozone layer, erosion due to acid rain, and damage from global climate change are caused by human activities. The ultimate effect of these changes on our carrying capacity is unknown.
54: Ecology of Individuals and Populations
Skills to Develop
• Describe how ecologists measure population size and density
• Describe three different patterns of population distribution
• Use life tables to calculate mortality rates
• Describe the three types of survivorship curves and relate them to specific populations
Populations are dynamic entities. Populations consist all of the species living within a specific area, and populations fluctuate based on a number of factors: seasonal and yearly changes in the environment, natural disasters such as forest fires and volcanic eruptions, and competition for resources between and within species. The statistical study of population dynamics, demography, uses a series of mathematical tools to investigate how populations respond to changes in their biotic and abiotic environments. Many of these tools were originally designed to study human populations. For example, life tables, which detail the life expectancy of individuals within a population, were initially developed by life insurance companies to set insurance rates. In fact, while the term “demographics” is commonly used when discussing humans, all living populations can be studied using this approach.
Population Size and Density
The study of any population usually begins by determining how many individuals of a particular species exist, and how closely associated they are with each other. Within a particular habitat, a population can be characterized by its population size (N), the total number of individuals, and its population density, the number of individuals within a specific area or volume. Population size and density are the two main characteristics used to describe and understand populations. For example, populations with more individuals may be more stable than smaller populations based on their genetic variability, and thus their potential to adapt to the environment. Alternatively, a member of a population with low population density (more spread out in the habitat), might have more difficulty finding a mate to reproduce compared to a population of higher density. As is shown in Figure $1$, smaller organisms tend to be more densely distributed than larger organisms.
Art Connection
As this graph shows, population density typically decreases with increasing body size. Why do you think this is the case?
Population Research Methods
The most accurate way to determine population size is to simply count all of the individuals within the habitat. However, this method is often not logistically or economically feasible, especially when studying large habitats. Thus, scientists usually study populations by sampling a representative portion of each habitat and using this data to make inferences about the habitat as a whole. A variety of methods can be used to sample populations to determine their size and density. For immobile organisms such as plants, or for very small and slow-moving organisms, a quadrat may be used (Figure $2$). A quadrat is a way of marking off square areas within a habitat, either by staking out an area with sticks and string, or by the use of a wood, plastic, or metal square placed on the ground. After setting the quadrats, researchers then count the number of individuals that lie within their boundaries. Multiple quadrat samples are performed throughout the habitat at several random locations. All of this data can then be used to estimate the population size and population density within the entire habitat. The number and size of quadrat samples depends on the type of organisms under study and other factors, including the density of the organism. For example, if sampling daffodils, a 1 m2 quadrat might be used whereas with giant redwoods, which are larger and live much further apart from each other, a larger quadrat of 100 m2 might be employed. This ensures that enough individuals of the species are counted to get an accurate sample that correlates with the habitat, including areas not sampled.
For mobile organisms, such as mammals, birds, or fish, a technique called mark and recapture is often used. This method involves marking a sample of captured animals in some way (such as tags, bands, paint, or other body markings), and then releasing them back into the environment to allow them to mix with the rest of the population; later, a new sample is collected, including some individuals that are marked (recaptures) and some individuals that are unmarked (Figure $3$).
Using the ratio of marked and unmarked individuals, scientists determine how many individuals are in the sample. From this, calculations are used to estimate the total population size. This method assumes that the larger the population, the lower the percentage of tagged organisms that will be recaptured since they will have mixed with more untagged individuals. For example, if 80 deer are captured, tagged, and released into the forest, and later 100 deer are captured and 20 of them are already marked, we can determine the population size (N) using the following equation:
$\frac{\text{number marked first catch} * \text{total number of second catch}} {\text{number marked second catch}} = N \nonumber$
Using our example, the population size would be estimated at 400.
$\frac{(80*100)} {20} = 400 \nonumber$
Therefore, there are an estimated 400 total individuals in the original population.
There are some limitations to the mark and recapture method. Some animals from the first catch may learn to avoid capture in the second round, thus inflating population estimates. Alternatively, animals may preferentially be retrapped (especially if a food reward is offered), resulting in an underestimate of population size. Also, some species may be harmed by the marking technique, reducing their survival. A variety of other techniques have been developed, including the electronic tracking of animals tagged with radio transmitters and the use of data from commercial fishing and trapping operations to estimate the size and health of populations and communities.
Species Distribution
In addition to measuring simple density, further information about a population can be obtained by looking at the distribution of the individuals. Species dispersion patterns (or distribution patterns) show the spatial relationship between members of a population within a habitat at a particular point in time. In other words, they show whether members of the species live close together or far apart, and what patterns are evident when they are spaced apart.
Individuals in a population can be more or less equally spaced apart, dispersed randomly with no predictable pattern, or clustered in groups. These are known as uniform, random, and clumped dispersion patterns, respectively (Figure $4$). Uniform dispersion is observed in plants that secrete substances inhibiting the growth of nearby individuals (such as the release of toxic chemicals by the sage plant Salvia leucophylla, a phenomenon called allelopathy) and in animals like the penguin that maintain a defined territory. An example of random dispersion occurs with dandelion and other plants that have wind-dispersed seeds that germinate wherever they happen to fall in a favorable environment. A clumped dispersion may be seen in plants that drop their seeds straight to the ground, such as oak trees, or animals that live in groups (schools of fish or herds of elephants). Clumped dispersions may also be a function of habitat heterogeneity. Thus, the dispersion of the individuals within a population provides more information about how they interact with each other than does a simple density measurement. Just as lower density species might have more difficulty finding a mate, solitary species with a random distribution might have a similar difficulty when compared to social species clumped together in groups.
Demography
While population size and density describe a population at one particular point in time, scientists must use demography to study the dynamics of a population. Demography is the statistical study of population changes over time: birth rates, death rates, and life expectancies. Each of these measures, especially birth rates, may be affected by the population characteristics described above. For example, a large population size results in a higher birth rate because more potentially reproductive individuals are present. In contrast, a large population size can also result in a higher death rate because of competition, disease, and the accumulation of waste. Similarly, a higher population density or a clumped dispersion pattern results in more potential reproductive encounters between individuals, which can increase birth rate. Lastly, a female-biased sex ratio (the ratio of males to females) or age structure (the proportion of population members at specific age ranges) composed of many individuals of reproductive age can increase birth rates.
In addition, the demographic characteristics of a population can influence how the population grows or declines over time. If birth and death rates are equal, the population remains stable. However, the population size will increase if birth rates exceed death rates; the population will decrease if birth rates are less than death rates. Life expectancy is another important factor; the length of time individuals remain in the population impacts local resources, reproduction, and the overall health of the population. These demographic characteristics are often displayed in the form of a life table.
Life Tables
Life tables provide important information about the life history of an organism. Life tables divide the population into age groups and often sexes, and show how long a member of that group is likely to live. They are modeled after actuarial tables used by the insurance industry for estimating human life expectancy. Life tables may include the probability of individuals dying before their next birthday (i.e., their mortality rate), the percentage of surviving individuals dying at a particular age interval, and their life expectancy at each interval. An example of a life table is shown in Table $1$ from a study of Dall mountain sheep, a species native to northwestern North America. Notice that the population is divided into age intervals (column A). The mortality rate (per 1000), shown in column D, is based on the number of individuals dying during the age interval (column B) divided by the number of individuals surviving at the beginning of the interval (Column C), multiplied by 1000.
$\text{mortality rate} = \frac{\text{number of individuals dying}} {\text{number of individuals surviving}} * 1000 \nonumber$
For example, between ages three and four, 12 individuals die out of the 776 that were remaining from the original 1000 sheep. This number is then multiplied by 1000 to get the mortality rate per thousand.
$\text{mortality rate} = \frac{12} {776} * 1000 \approx 15.5 \nonumber$
As can be seen from the mortality rate data (column D), a high death rate occurred when the sheep were between 6 and 12 months old, and then increased even more from 8 to 12 years old, after which there were few survivors. The data indicate that if a sheep in this population were to survive to age one, it could be expected to live another 7.7 years on average, as shown by the life expectancy numbers in column E.
Table $1$: Life Table of Dall Mountain Sheep1
Age interval (years) Number dying in age interval out of 1000 born Number surviving at beginning of age interval out of 1000 born Mortality rate per 1000 alive at beginning of age interval Life expectancy or mean lifetime remaining to those attaining age interval
0-0.5 54 1000 54.0 7.06
0.5-1 145 946 153.3 --
1-2 12 801 15.0 7.7
2-3 13 789 16.5 6.8
3-4 12 776 15.5 5.9
4-5 30 764 39.3 5.0
5-6 46 734 62.7 4.2
6-7 48 688 69.8 3.4
7-8 69 640 107.8 2.6
8-9 132 571 231.2 1.9
9-10 187 439 426.0 1.3
10-11 156 252 619.0 0.9
11-12 90 96 937.5 0.6
12-13 3 6 500.0 1.2
13-14 3 3 1000 0.7
Survivorship Curves
Another tool used by population ecologists is a survivorship curve, which is a graph of the number of individuals surviving at each age interval plotted versus time (usually with data compiled from a life table). These curves allow us to compare the life histories of different populations (Figure $5$). Humans and most primates exhibit a Type I survivorship curve because a high percentage of offspring survive their early and middle years—death occurs predominantly in older individuals. These types of species usually have small numbers of offspring at one time, and they give a high amount of parental care to them to ensure their survival. Birds are an example of an intermediate or Type II survivorship curve because birds die more or less equally at each age interval. These organisms also may have relatively few offspring and provide significant parental care. Trees, marine invertebrates, and most fishes exhibit a Type III survivorship curve because very few of these organisms survive their younger years; however, those that make it to an old age are more likely to survive for a relatively long period of time. Organisms in this category usually have a very large number of offspring, but once they are born, little parental care is provided. Thus these offspring are “on their own” and vulnerable to predation, but their sheer numbers assure the survival of enough individuals to perpetuate the species.
Summary
Populations are individuals of a species that live in a particular habitat. Ecologists measure characteristics of populations: size, density, dispersion pattern, age structure, and sex ratio. Life tables are useful to calculate life expectancies of individual population members. Survivorship curves show the number of individuals surviving at each age interval plotted versus time.
Art Connections
Figure $1$: As this graph shows, population density typically decreases with increasing body size. Why do you think this is the case?
Answer
Smaller animals require less food and other resources, so the environment can support more of them.
Footnotes
1. 1 Data Adapted from Edward S. Deevey, Jr., “Life Tables for Natural Populations of Animals,” The Quarterly Review of Biology 22, no. 4 (December 1947): 283-314.
Glossary
demography
statistical study of changes in populations over time
life table
table showing the life expectancy of a population member based on its age
mark and recapture
technique used to determine population size in mobile organisms
mortality rate
proportion of population surviving to the beginning of an age interval that die during the age interval
population density
number of population members divided by the area or volume being measured
population size (N)
number of population members in a habitat at the same time
quadrat
square made of various materials used to determine population size and density in slow moving or stationary organisms
species dispersion pattern
(also, species distribution pattern) spatial location of individuals of a given species within a habitat at a particular point in time
survivorship curve
graph of the number of surviving population members versus the relative age of the member | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/54%3A_Ecology_of_Individuals_and_Populations/54.01%3A_The_Environmental_Challenges/54.1.01%3A_Prelude_to_Population_and_Community_Ecology.txt |
Skills to Develop
• Describe how life history patterns are influenced by natural selection
• Explain different life history patterns and how different reproductive strategies affect species’ survival
A species’ life history describes the series of events over its lifetime, such as how resources are allocated for growth, maintenance, and reproduction. Life history traits affect the life table of an organism. A species’ life history is genetically determined and shaped by the environment and natural selection.
Life History Patterns and Energy Budgets
Energy is required by all living organisms for their growth, maintenance, and reproduction; at the same time, energy is often a major limiting factor in determining an organism’s survival. Plants, for example, acquire energy from the sun via photosynthesis, but must expend this energy to grow, maintain health, and produce energy-rich seeds to produce the next generation. Animals have the additional burden of using some of their energy reserves to acquire food. Furthermore, some animals must expend energy caring for their offspring. Thus, all species have an energy budget: they must balance energy intake with their use of energy for metabolism, reproduction, parental care, and energy storage (such as bears building up body fat for winter hibernation).
Parental Care and Fecundity
Fecundity is the potential reproductive capacity of an individual within a population. In other words, fecundity describes how many offspring could ideally be produced if an individual has as many offspring as possible, repeating the reproductive cycle as soon as possible after the birth of the offspring. In animals, fecundity is inversely related to the amount of parental care given to an individual offspring. Species, such as many marine invertebrates, that produce many offspring usually provide little if any care for the offspring (they would not have the energy or the ability to do so anyway). Most of their energy budget is used to produce many tiny offspring. Animals with this strategy are often self-sufficient at a very early age. This is because of the energy tradeoff these organisms have made to maximize their evolutionary fitness. Because their energy is used for producing offspring instead of parental care, it makes sense that these offspring have some ability to be able to move within their environment and find food and perhaps shelter. Even with these abilities, their small size makes them extremely vulnerable to predation, so the production of many offspring allows enough of them to survive to maintain the species.
Animal species that have few offspring during a reproductive event usually give extensive parental care, devoting much of their energy budget to these activities, sometimes at the expense of their own health. This is the case with many mammals, such as humans, kangaroos, and pandas. The offspring of these species are relatively helpless at birth and need to develop before they achieve self-sufficiency.
Plants with low fecundity produce few energy-rich seeds (such as coconuts and chestnuts) with each having a good chance to germinate into a new organism; plants with high fecundity usually have many small, energy-poor seeds (like orchids) that have a relatively poor chance of surviving. Although it may seem that coconuts and chestnuts have a better chance of surviving, the energy tradeoff of the orchid is also very effective. It is a matter of where the energy is used, for large numbers of seeds or for fewer seeds with more energy.
Early versus Late Reproduction
The timing of reproduction in a life history also affects species survival. Organisms that reproduce at an early age have a greater chance of producing offspring, but this is usually at the expense of their growth and the maintenance of their health. Conversely, organisms that start reproducing later in life often have greater fecundity or are better able to provide parental care, but they risk that they will not survive to reproductive age. Examples of this can be seen in fishes. Small fish like guppies use their energy to reproduce rapidly, but never attain the size that would give them defense against some predators. Larger fish, like the bluegill or shark, use their energy to attain a large size, but do so with the risk that they will die before they can reproduce or at least reproduce to their maximum. These different energy strategies and tradeoffs are key to understanding the evolution of each species as it maximizes its fitness and fills its niche. In terms of energy budgeting, some species “blow it all” and use up most of their energy reserves to reproduce early before they die. Other species delay having reproduction to become stronger, more experienced individuals and to make sure that they are strong enough to provide parental care if necessary.
Single versus Multiple Reproductive Events
Some life history traits, such as fecundity, timing of reproduction, and parental care, can be grouped together into general strategies that are used by multiple species. Semelparity occurs when a species reproduces only once during its lifetime and then dies. Such species use most of their resource budget during a single reproductive event, sacrificing their health to the point that they do not survive. Examples of semelparity are bamboo, which flowers once and then dies, and the Chinook salmon (Figure \(1\)a), which uses most of its energy reserves to migrate from the ocean to its freshwater nesting area, where it reproduces and then dies. Scientists have posited alternate explanations for the evolutionary advantage of the Chinook’s post-reproduction death: a programmed suicide caused by a massive release of corticosteroid hormones, presumably so the parents can become food for the offspring, or simple exhaustion caused by the energy demands of reproduction; these are still being debated.
Iteroparity describes species that reproduce repeatedly during their lives. Some animals are able to mate only once per year, but survive multiple mating seasons. The pronghorn antelope is an example of an animal that goes into a seasonal estrus cycle (“heat”): a hormonally induced physiological condition preparing the body for successful mating (Figure \(1\)b). Females of these species mate only during the estrus phase of the cycle. A different pattern is observed in primates, including humans and chimpanzees, which may attempt reproduction at any time during their reproductive years, even though their menstrual cycles make pregnancy likely only a few days per month during ovulation (Figure \(1\)c).
Evolution Connection: Energy Budgets, Reproductive Costs, and Sexual Selection in Drosophila
Research into how animals allocate their energy resources for growth, maintenance, and reproduction has used a variety of experimental animal models. Some of this work has been done using the common fruit fly, Drosophila melanogaster. Studies have shown that not only does reproduction have a cost as far as how long male fruit flies live, but also fruit flies that have already mated several times have limited sperm remaining for reproduction. Fruit flies maximize their last chances at reproduction by selecting optimal mates.
In a 1981 study, male fruit flies were placed in enclosures with either virgin or inseminated females. The males that mated with virgin females had shorter life spans than those in contact with the same number of inseminated females with which they were unable to mate. This effect occurred regardless of how large (indicative of their age) the males were. Thus, males that did not mate lived longer, allowing them more opportunities to find mates in the future.
More recent studies, performed in 2006, show how males select the female with which they will mate and how this is affected by previous matings (Figure \(2\)).1 Males were allowed to select between smaller and larger females. Findings showed that larger females had greater fecundity, producing twice as many offspring per mating as the smaller females did. Males that had previously mated, and thus had lower supplies of sperm, were termed “resource-depleted,” while males that had not mated were termed “non-resource-depleted.” The study showed that although non-resource-depleted males preferentially mated with larger females, this selection of partners was more pronounced in the resource-depleted males. Thus, males with depleted sperm supplies, which were limited in the number of times that they could mate before they replenished their sperm supply, selected larger, more fecund females, thus maximizing their chances for offspring. This study was one of the first to show that the physiological state of the male affected its mating behavior in a way that clearly maximizes its use of limited reproductive resources.
These studies demonstrate two ways in which the energy budget is a factor in reproduction. First, energy expended on mating may reduce an animal’s lifespan, but by this time they have already reproduced, so in the context of natural selection this early death is not of much evolutionary importance. Second, when resources such as sperm (and the energy needed to replenish it) are low, an organism’s behavior can change to give them the best chance of passing their genes on to the next generation. These changes in behavior, so important to evolution, are studied in a discipline known as behavioral biology, or ethology, at the interface between population biology and psychology.
Summary
All species have evolved a pattern of living, called a life history strategy, in which they partition energy for growth, maintenance, and reproduction. These patterns evolve through natural selection; they allow species to adapt to their environment to obtain the resources they need to successfully reproduce. There is an inverse relationship between fecundity and parental care. A species may reproduce early in life to ensure surviving to a reproductive age or reproduce later in life to become larger and healthier and better able to give parental care. A species may reproduce once (semelparity) or many times (iteroparity) in its life.
Footnotes
1. 1 Adapted from Phillip G. Byrne and William R. Rice, “Evidence for adaptive male mate choice in the fruit fly Drosophila melanogaster,” Proc Biol Sci. 273, no. 1589 (2006): 917-922, doi: 10.1098/rspb.2005.3372.
Glossary
energy budget
allocation of energy resources for body maintenance, reproduction, and parental care
fecundity
potential reproductive capacity of an individual
iteroparity
life history strategy characterized by multiple reproductive events during the lifetime of a species
life history
inherited pattern of resource allocation under the influence of natural selection and other evolutionary forces
semelparity
life history strategy characterized by a single reproductive event followed by death | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/54%3A_Ecology_of_Individuals_and_Populations/54.04%3A_Life_History_and_the_Cost_of_Reproduction.txt |
Skills to Develop
• Explain the characteristics of and differences between exponential and logistic growth patterns
• Give examples of exponential and logistic growth in natural populations
• Describe how natural selection and environmental adaptation led to the evolution of particular life history patterns
Although life histories describe the way many characteristics of a population (such as their age structure) change over time in a general way, population ecologists make use of a variety of methods to model population dynamics mathematically. These more precise models can then be used to accurately describe changes occurring in a population and better predict future changes. Certain models that have been accepted for decades are now being modified or even abandoned due to their lack of predictive ability, and scholars strive to create effective new models.
Exponential Growth
Charles Darwin, in his theory of natural selection, was greatly influenced by the English clergyman Thomas Malthus. Malthus published a book in 1798 stating that populations with unlimited natural resources grow very rapidly, and then population growth decreases as resources become depleted. This accelerating pattern of increasing population size is called exponential growth.
The best example of exponential growth is seen in bacteria. Bacteria are prokaryotes that reproduce by prokaryotic fission. This division takes about an hour for many bacterial species. If 1000 bacteria are placed in a large flask with an unlimited supply of nutrients (so the nutrients will not become depleted), after an hour, there is one round of division and each organism divides, resulting in 2000 organisms—an increase of 1000. In another hour, each of the 2000 organisms will double, producing 4000, an increase of 2000 organisms. After the third hour, there should be 8000 bacteria in the flask, an increase of 4000 organisms. The important concept of exponential growth is that the population growth rate—the number of organisms added in each reproductive generation—is accelerating; that is, it is increasing at a greater and greater rate. After 1 day and 24 of these cycles, the population would have increased from 1000 to more than 16 billion. When the population size, N, is plotted over time, a J-shaped growth curve is produced (Figure $1$).
The bacteria example is not representative of the real world where resources are limited. Furthermore, some bacteria will die during the experiment and thus not reproduce, lowering the growth rate. Therefore, when calculating the growth rate of a population, the death rate (D) (number organisms that die during a particular time interval) is subtracted from the birth rate (B) (number organisms that are born during that interval). This is shown in the following formula:
$\frac{\Delta N\;(\text{change in number})} {\Delta T\;(\text{change in time})} = B\;(\text{birth rate}) - D\;(\text{death rate}) \nonumber$
The birth rate is usually expressed on a per capita (for each individual) basis. Thus, B (birth rate) = bN (the per capita birth rate “b” multiplied by the number of individuals “N”) and D (death rate) =dN (the per capita death rate “d” multiplied by the number of individuals “N”). Additionally, ecologists are interested in the population at a particular point in time, an infinitely small time interval. For this reason, the terminology of differential calculus is used to obtain the “instantaneous” growth rate, replacing the change in number and time with an instant-specific measurement of number and time.
$\frac{d N} {d T} = bN - dN = (b - d)N \nonumber$
Notice that the “d” associated with the first term refers to the derivative (as the term is used in calculus) and is different from the death rate, also called “$d$.” The difference between birth and death rates is further simplified by substituting the term “r” (intrinsic rate of increase) for the relationship between birth and death rates:
$\frac{d N} {d T} = rN \nonumber$
The value “$r$” can be positive, meaning the population is increasing in size; or negative, meaning the population is decreasing in size; or zero, where the population’s size is unchanging, a condition known as zero population growth. A further refinement of the formula recognizes that different species have inherent differences in their intrinsic rate of increase (often thought of as the potential for reproduction), even under ideal conditions. Obviously, a bacterium can reproduce more rapidly and have a higher intrinsic rate of growth than a human. The maximal growth rate for a species is its biotic potential, or $r_{max}$, thus changing the equation to:
$\frac{d N} {d T} = r_\text{max}N \nonumber$
Logistic Growth
Exponential growth is possible only when infinite natural resources are available; this is not the case in the real world. Charles Darwin recognized this fact in his description of the “struggle for existence,” which states that individuals will compete (with members of their own or other species) for limited resources. The successful ones will survive to pass on their own characteristics and traits (which we know now are transferred by genes) to the next generation at a greater rate (natural selection). To model the reality of limited resources, population ecologists developed the logistic growth model.
Carrying Capacity and the Logistic Model
In the real world, with its limited resources, exponential growth cannot continue indefinitely. Exponential growth may occur in environments where there are few individuals and plentiful resources, but when the number of individuals gets large enough, resources will be depleted, slowing the growth rate. Eventually, the growth rate will plateau or level off (Figure $1$). This population size, which represents the maximum population size that a particular environment can support, is called the carrying capacity, or $K$.
The formula we use to calculate logistic growth adds the carrying capacity as a moderating force in the growth rate. The expression “$K – N$” is indicative of how many individuals may be added to a population at a given stage, and “$K – N$” divided by “$K$” is the fraction of the carrying capacity available for further growth. Thus, the exponential growth model is restricted by this factor to generate the logistic growth equation:
\begin{align} \dfrac{d N} {d T} &= r_\text{max} \frac{d N} {d T} \nonumber \[5pt] &= r_\text{max}N \frac{(K-N)} {K}\nonumber \end{align} \nonumber
Notice that when $N$ is very small, $(K-N)/K$ becomes close to $K/K$ or $1$, and the right side of the equation reduces to $r_{max}N$, which means the population is growing exponentially and is not influenced by carrying capacity. On the other hand, when $N$ is large, $(K-N)/K$ come close to zero, which means that population growth will be slowed greatly or even stopped. Thus, population growth is greatly slowed in large populations by the carrying capacity $K$. This model also allows for the population of a negative population growth, or a population decline. This occurs when the number of individuals in the population exceeds the carrying capacity (because the value of $(K-N)/K$ is negative).
A graph of this equation yields an S-shaped curve (Figure $1$), and it is a more realistic model of population growth than exponential growth. There are three different sections to an S-shaped curve. Initially, growth is exponential because there are few individuals and ample resources available. Then, as resources begin to become limited, the growth rate decreases. Finally, growth levels off at the carrying capacity of the environment, with little change in population size over time.
Role of Intraspecific Competition
The logistic model assumes that every individual within a population will have equal access to resources and, thus, an equal chance for survival. For plants, the amount of water, sunlight, nutrients, and the space to grow are the important resources, whereas in animals, important resources include food, water, shelter, nesting space, and mates.
In the real world, phenotypic variation among individuals within a population means that some individuals will be better adapted to their environment than others. The resulting competition between population members of the same species for resources is termed intraspecific competition (intra- = “within”; -specific = “species”). Intraspecific competition for resources may not affect populations that are well below their carrying capacity—resources are plentiful and all individuals can obtain what they need. However, as population size increases, this competition intensifies. In addition, the accumulation of waste products can reduce an environment’s carrying capacity.
Examples of Logistic Growth
Yeast, a microscopic fungus used to make bread and alcoholic beverages, exhibits the classical S-shaped curve when grown in a test tube (Figure $2$a). Its growth levels off as the population depletes the nutrients that are necessary for its growth. In the real world, however, there are variations to this idealized curve. Examples in wild populations include sheep and harbor seals (Figure $2$b). In both examples, the population size exceeds the carrying capacity for short periods of time and then falls below the carrying capacity afterwards. This fluctuation in population size continues to occur as the population oscillates around its carrying capacity. Still, even with this oscillation, the logistic model is confirmed.
Art Connection
If the major food source of the seals declines due to pollution or overfishing, which of the following would likely occur?
1. The carrying capacity of seals would decrease, as would the seal population.
2. The carrying capacity of seals would decrease, but the seal population would remain the same.
3. The number of seal deaths would increase but the number of births would also increase, so the population size would remain the same.
4. The carrying capacity of seals would remain the same, but the population of seals would decrease.
Summary
Populations with unlimited resources grow exponentially, with an accelerating growth rate. When resources become limiting, populations follow a logistic growth curve. The population of a species will level off at the carrying capacity of its environment.
Art Connections
Figure $2$b If the major food source of the seals declines due to pollution or overfishing, which of the following would likely occur?
1. The carrying capacity of seals would decrease, as would the seal population.
2. The carrying capacity of seals would decrease, but the seal population would remain the same.
3. The number of seal deaths would increase but the number of births would also increase, so the population size would remain the same.
4. The carrying capacity of seals would remain the same, but the population of seals would decrease.
Answer
A
Glossary
biotic potential (rmax)
maximal potential growth rate of a species
birth rate (B)
number of births within a population at a specific point in time
carrying capacity (K)
number of individuals of a species that can be supported by the limited resources of a habitat
death rate (D)
number of deaths within a population at a specific point in time
exponential growth
accelerating growth pattern seen in species under conditions where resources are not limiting
intraspecific competition
competition between members of the same species
J-shaped growth curve
shape of an exponential growth curve
logistic growth
leveling off of exponential growth due to limiting resources
population growth rate
number of organisms added in each reproductive generation
S-shaped growth curve
shape of a logistic growth curve
zero population growth
steady population size where birth rates and death rates are equal | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/54%3A_Ecology_of_Individuals_and_Populations/54.05%3A_Environmental_Limits_to_Population_Growth.txt |
Skills to Develop
• Give examples of how the carrying capacity of a habitat may change
• Compare and contrast density-dependent growth regulation and density-independent growth regulation, giving examples
• Give examples of exponential and logistic growth in wild animal populations
• Describe how natural selection and environmental adaptation leads to the evolution of particular life-history patterns
The logistic model of population growth, while valid in many natural populations and a useful model, is a simplification of real-world population dynamics. Implicit in the model is that the carrying capacity of the environment does not change, which is not the case. The carrying capacity varies annually: for example, some summers are hot and dry whereas others are cold and wet. In many areas, the carrying capacity during the winter is much lower than it is during the summer. Also, natural events such as earthquakes, volcanoes, and fires can alter an environment and hence its carrying capacity. Additionally, populations do not usually exist in isolation. They engage in interspecific competition: that is, they share the environment with other species, competing with them for the same resources. These factors are also important to understanding how a specific population will grow.
Nature regulates population growth in a variety of ways. These are grouped into density-dependent factors, in which the density of the population at a given time affects growth rate and mortality, and density-independent factors, which influence mortality in a population regardless of population density. Note that in the former, the effect of the factor on the population depends on the density of the population at onset. Conservation biologists want to understand both types because this helps them manage populations and prevent extinction or overpopulation.
Density-dependent Regulation
Most density-dependent factors are biological in nature (biotic), and include predation, inter- and intraspecific competition, accumulation of waste, and diseases such as those caused by parasites. Usually, the denser a population is, the greater its mortality rate. For example, during intra- and interspecific competition, the reproductive rates of the individuals will usually be lower, reducing their population’s rate of growth. In addition, low prey density increases the mortality of its predator because it has more difficulty locating its food source.
An example of density-dependent regulation is shown in Figure \(1\) with results from a study focusing on the giant intestinal roundworm (Ascaris lumbricoides), a parasite of humans and other mammals.1 Denser populations of the parasite exhibited lower fecundity: they contained fewer eggs. One possible explanation for this is that females would be smaller in more dense populations (due to limited resources) and that smaller females would have fewer eggs. This hypothesis was tested and disproved in a 2009 study which showed that female weight had no influence.2 The actual cause of the density-dependence of fecundity in this organism is still unclear and awaiting further investigation.
Density-independent Regulation and Interaction with Density-dependent Factors
Many factors, typically physical or chemical in nature (abiotic), influence the mortality of a population regardless of its density, including weather, natural disasters, and pollution. An individual deer may be killed in a forest fire regardless of how many deer happen to be in that area. Its chances of survival are the same whether the population density is high or low. The same holds true for cold winter weather.
In real-life situations, population regulation is very complicated and density-dependent and independent factors can interact. A dense population that is reduced in a density-independent manner by some environmental factor(s) will be able to recover differently than a sparse population. For example, a population of deer affected by a harsh winter will recover faster if there are more deer remaining to reproduce.
Evolution Connection: Why Did the Woolly Mammoth Go Extinct?
It's easy to get lost in the discussion of dinosaurs and theories about why they went extinct 65 million years ago. Was it due to a meteor slamming into Earth near the coast of modern-day Mexico, or was it from some long-term weather cycle that is not yet understood? One hypothesis that will never be proposed is that humans had something to do with it. Mammals were small, insignificant creatures of the forest 65 million years ago, and no humans existed.
Woolly mammoths, however, began to go extinct about 10,000 years ago, when they shared the Earth with humans who were no different anatomically than humans today (Figure \(2\)). Mammoths survived in isolated island populations as recently as 1700 BC. We know a lot about these animals from carcasses found frozen in the ice of Siberia and other regions of the north. Scientists have sequenced at least 50 percent of its genome and believe mammoths are between 98 and 99 percent identical to modern elephants.
It is commonly thought that climate change and human hunting led to their extinction. A 2008 study estimated that climate change reduced the mammoth’s range from 3,000,000 square miles 42,000 years ago to 310,000 square miles 6,000 years ago.4 It is also well documented that humans hunted these animals. A 2012 study showed that no single factor was exclusively responsible for the extinction of these magnificent creatures.5 In addition to human hunting, climate change, and reduction of habitat, these scientists demonstrated another important factor in the mammoth’s extinction was the migration of humans across the Bering Strait to North America during the last ice age 20,000 years ago.
The maintenance of stable populations was and is very complex, with many interacting factors determining the outcome. It is important to remember that humans are also part of nature. Once we contributed to a species’ decline using primitive hunting technology only.
Life Histories of K-selected and r-selected Species
While reproductive strategies play a key role in life histories, they do not account for important factors like limited resources and competition. The regulation of population growth by these factors can be used to introduce a classical concept in population biology, that of K-selected versus r-selected species.
Early Theories about Life History: K-selected and r-selected Species
By the second half of the twentieth century, the concept of K- and r-selected species was used extensively and successfully to study populations. The concept relates not only reproductive strategies, but also to a species’ habitat and behavior, especially in the way that they obtain resources and care for their young. It includes length of life and survivorship factors as well. For this analysis, population biologists have grouped species into the two large categories—K-selected and r-selected—although they are really two ends of a continuum.
K-selected species are species selected by stable, predictable environments. Populations of K-selected species tend to exist close to their carrying capacity (hence the term K-selected) where intraspecific competition is high. These species have few, large offspring, a long gestation period, and often give long-term care to their offspring (Table \(1\)). While larger in size when born, the offspring are relatively helpless and immature at birth. By the time they reach adulthood, they must develop skills to compete for natural resources. In plants, scientists think of parental care more broadly: how long fruit takes to develop or how long it remains on the plant are determining factors in the time to the next reproductive event. Examples of K-selected species are primates including humans), elephants, and plants such as oak trees (Figure \(3\)a).
Oak trees grow very slowly and take, on average, 20 years to produce their first seeds, known as acorns. As many as 50,000 acorns can be produced by an individual tree, but the germination rate is low as many of these rot or are eaten by animals such as squirrels. In some years, oaks may produce an exceptionally large number of acorns, and these years may be on a two- or three-year cycle depending on the species of oak (r-selection).
As oak trees grow to a large size and for many years before they begin to produce acorns, they devote a large percentage of their energy budget to growth and maintenance. The tree’s height and size allow it to dominate other plants in the competition for sunlight, the oak’s primary energy resource. Furthermore, when it does reproduce, the oak produces large, energy-rich seeds that use their energy reserve to become quickly established (K-selection).
In contrast, r-selected species have a large number of small offspring (hence their r designation (Table \(1\)). This strategy is often employed in unpredictable or changing environments. Animals that are r-selected do not give long-term parental care and the offspring are relatively mature and self-sufficient at birth. Examples of r-selected species are marine invertebrates, such as jellyfish, and plants, such as the dandelion (Figure \(3\)b). Dandelions have small seeds that are wind dispersed long distances. Many seeds are produced simultaneously to ensure that at least some of them reach a hospitable environment. Seeds that land in inhospitable environments have little chance for survival since their seeds are low in energy content. Note that survival is not necessarily a function of energy stored in the seed itself.
Table \(1\): Characteristics of K-selected and r-selected species
Characteristics of K-selected species Characteristics of r-selected species
Mature late Mature early
Greater longevity Lower longevity
Increased parental care Decreased parental care
Increased competition Decreased competition
Fewer offspring More offspring
Larger offspring Smaller offspring
Modern Theories of Life History
The r- and K-selection theory, although accepted for decades and used for much groundbreaking research, has now been reconsidered, and many population biologists have abandoned or modified it. Over the years, several studies attempted to confirm the theory, but these attempts have largely failed. Many species were identified that did not follow the theory’s predictions. Furthermore, the theory ignored the age-specific mortality of the populations which scientists now know is very important. New demographic-based models of life history evolution have been developed which incorporate many ecological concepts included in r- and K-selection theory as well as population age structure and mortality factors.
Summary
Populations are regulated by a variety of density-dependent and density-independent factors. Species are divided into two categories based on a variety of features of their life history patterns: r-selected species, which have large numbers of offspring, and K-selected species, which have few offspring. The r- and K-selection theory has fallen out of use; however, many of its key features are still used in newer, demographically-based models of population dynamics.
Footnotes
1. 1 N.A. Croll et al., “The Population Biology and Control of Ascaris lumbricoides in a Rural Community in Iran.” Transactions of the Royal Society of Tropical Medicine and Hygiene 76, no. 2 (1982): 187-197, doi:10.1016/0035-9203(82)90272-3.
2. 2 Martin Walker et al., “Density-Dependent Effects on the Weight of Female Ascaris lumbricoides Infections of Humans and its Impact on Patterns of Egg Production.” Parasites & Vectors 2, no. 11 (February 2009), doi:10.1186/1756-3305-2-11.
3. 3 N.A. Croll et al., “The Population Biology and Control of Ascaris lumbricoides in a Rural Community in Iran.” Transactions of the Royal Society of Tropical Medicine and Hygiene 76, no. 2 (1982): 187-197, doi:10.1016/0035-9203(82)90272-3.
4. 4 David Nogués-Bravo et al., “Climate Change, Humans, and the Extinction of the Woolly Mammoth.” PLoS Biol 6 (April 2008): e79, doi:10.1371/journal.pbio.0060079.
5. 5 G.M. MacDonald et al., “Pattern of Extinction of the Woolly Mammoth in Beringia.” Nature Communications 3, no. 893 (June 2012), doi:10.1038/ncomms1881.
Glossary
demographic-based population model
modern model of population dynamics incorporating many features of the r- and K-selection theory
density-dependent regulation
regulation of population that is influenced by population density, such as crowding effects; usually involves biotic factors
density-independent regulation
regulation of populations by factors that operate independent of population density, such as forest fires and volcanic eruptions; usually involves abiotic factors
interspecific competition
competition between species for resources in a shared habitat or environment
K-selected species
species suited to stable environments that produce a few, relatively large offspring and provide parental care
r-selected species
species suited to changing environments that produce many offspring and provide little or no parental care | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/54%3A_Ecology_of_Individuals_and_Populations/54.06%3A_Factors_that_Regulate_Populations.txt |
Skills to Develop
• Discuss how human population growth can be exponential
• Explain how humans have expanded the carrying capacity of their habitat
• Relate population growth and age structure to the level of economic development in different countries
• Discuss the long-term implications of unchecked human population growth
Concepts of animal population dynamics can be applied to human population growth. Humans are not unique in their ability to alter their environment. For example, beaver dams alter the stream environment where they are built. Humans, however, have the ability to alter their environment to increase its carrying capacity sometimes to the detriment of other species (e.g., via artificial selection for crops that have a higher yield). Earth’s human population is growing rapidly, to the extent that some worry about the ability of the earth’s environment to sustain this population, as long-term exponential growth carries the potential risks of famine, disease, and large-scale death.
Although humans have increased the carrying capacity of their environment, the technologies used to achieve this transformation have caused unprecedented changes to Earth’s environment, altering ecosystems to the point where some may be in danger of collapse. The depletion of the ozone layer, erosion due to acid rain, and damage from global climate change are caused by human activities. The ultimate effect of these changes on our carrying capacity is unknown. As some point out, it is likely that the negative effects of increasing carrying capacity will outweigh the positive ones—the carrying capacity of the world for human beings might actually decrease.
The world’s human population is currently experiencing exponential growth even though human reproduction is far below its biotic potential (Figure \(1\)). To reach its biotic potential, all females would have to become pregnant every nine months or so during their reproductive years. Also, resources would have to be such that the environment would support such growth. Neither of these two conditions exists. In spite of this fact, human population is still growing exponentially.
A consequence of exponential human population growth is the time that it takes to add a particular number of humans to the Earth is becoming shorter. Figure \(2\) shows that 123 years were necessary to add 1 billion humans in 1930, but it only took 24 years to add two billion people between 1975 and 1999. As already discussed, at some point it would appear that our ability to increase our carrying capacity indefinitely on a finite world is uncertain. Without new technological advances, the human growth rate has been predicted to slow in the coming decades. However, the population will still be increasing and the threat of overpopulation remains.
Link to Learning
Click through this interactive view of how human populations have changed over time.
Overcoming Density-Dependent Regulation
Humans are unique in their ability to alter their environment with the conscious purpose of increasing its carrying capacity. This ability is a major factor responsible for human population growth and a way of overcoming density-dependent growth regulation. Much of this ability is related to human intelligence, society, and communication. Humans can construct shelter to protect them from the elements and have developed agriculture and domesticated animals to increase their food supplies. In addition, humans use language to communicate this technology to new generations, allowing them to improve upon previous accomplishments.
Other factors in human population growth are migration and public health. Humans originated in Africa, but have since migrated to nearly all inhabitable land on the Earth. Public health, sanitation, and the use of antibiotics and vaccines have decreased the ability of infectious disease to limit human population growth. In the past, diseases such as the bubonic plaque of the fourteenth century killed between 30 and 60 percent of Europe’s population and reduced the overall world population by as many as 100 million people. Today, the threat of infectious disease, while not gone, is certainly less severe. According to the World Health Organization, global death from infectious disease declined from 16.4 million in 1993 to 14.7 million in 1992. To compare to some of the epidemics of the past, the percentage of the world's population killed between 1993 and 2002 decreased from 0.30 percent of the world's population to 0.24 percent. Thus, it appears that the influence of infectious disease on human population growth is becoming less significant.
Age Structure, Population Growth, and Economic Development
The age structure of a population is an important factor in population dynamics. Age structure is the proportion of a population at different age ranges. Age structure allows better prediction of population growth, plus the ability to associate this growth with the level of economic development in the region. Countries with rapid growth have a pyramidal shape in their age structure diagrams, showing a preponderance of younger individuals, many of whom are of reproductive age or will be soon (Figure \(3\)). This pattern is most often observed in underdeveloped countries where individuals do not live to old age because of less-than-optimal living conditions. Age structures of areas with slow growth, including developed countries such as the United States, still have a pyramidal structure, but with many fewer young and reproductive-aged individuals and a greater proportion of older individuals. Other developed countries, such as Italy, have zero population growth. The age structure of these populations is more conical, with an even greater percentage of middle-aged and older individuals. The actual growth rates in different countries are shown in Figure \(4\), with the highest rates tending to be in the less economically developed countries of Africa and Asia.
Art Connection
Age structure diagrams for rapidly growing, slow growing and stable populations are shown in stages 1 through 3. What type of population change do you think stage 4 represents?
Long-Term Consequences of Exponential Human Population Growth
Many dire predictions have been made about the world’s population leading to a major crisis called the “population explosion.” In the 1968 book The Population Bomb, biologist Dr. Paul R. Ehrlich wrote, “The battle to feed all of humanity is over. In the 1970s hundreds of millions of people will starve to death in spite of any crash programs embarked upon now. At this late date nothing can prevent a substantial increase in the world death rate.”1 While many critics view this statement as an exaggeration, the laws of exponential population growth are still in effect, and unchecked human population growth cannot continue indefinitely.
Efforts to control population growth led to the one-child policy in China, which used to include more severe consequences, but now imposes fines on urban couples who have more than one child. Due to the fact that some couples wish to have a male heir, many Chinese couples continue to have more than one child. The policy itself, its social impacts, and the effectiveness of limiting overall population growth are controversial. In spite of population control policies, the human population continues to grow. At some point the food supply may run out because of the subsequent need to produce more and more food to feed our population. The United Nations estimates that future world population growth may vary from 6 billion (a decrease) to 16 billion people by the year 2100. There is no way to know whether human population growth will moderate to the point where the crisis described by Dr. Ehrlich will be averted.
Another result of population growth is the endangerment of the natural environment. Many countries have attempted to reduce the human impact on climate change by reducing their emission of the greenhouse gas carbon dioxide. However, these treaties have not been ratified by every country, and many underdeveloped countries trying to improve their economic condition may be less likely to agree with such provisions if it means slower economic development. Furthermore, the role of human activity in causing climate change has become a hotly debated socio-political issue in some developed countries, including the United States. Thus, we enter the future with considerable uncertainty about our ability to curb human population growth and protect our environment.
Link to Learning
Visit this website and select “Launch movie” for an animation discussing the global impacts of human population growth.
Summary
The world’s human population is growing at an exponential rate. Humans have increased the world’s carrying capacity through migration, agriculture, medical advances, and communication. The age structure of a population allows us to predict population growth. Unchecked human population growth could have dire long-term effects on our environment.
Art Connections
Figure \(3\): Age structure diagrams for rapidly growing, slow growing and stable populations are shown in stages 1 through 3. What type of population change do you think stage 4 represents?
Answer
Stage 4 represents a population that is decreasing.
Footnotes
1. 1 Paul R. Erlich, prologue to The Population Bomb, (1968; repr., New York: Ballantine, 1970).
Glossary
age structure
proportion of population members at specific age ranges
one-child policy
China’s policy to limit population growth by limiting urban couples to have only one child or face the penalty of a fine | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/54%3A_Ecology_of_Individuals_and_Populations/54.07%3A_Human_Population_Growth.txt |
Skills to Develop
• Define biogeography
• List and describe abiotic factors that affect the global distribution of plant and animal species
• Compare the impact of abiotic forces on aquatic and terrestrial environments
• Summarize the affect of abiotic factors on net primary productivity
Many forces influence the communities of living organisms present in different parts of the biosphere (all of the parts of Earth inhabited by life). The biosphere extends into the atmosphere (several kilometers above Earth) and into the depths of the oceans. Despite its apparent vastness to an individual human, the biosphere occupies only a minute space when compared to the known universe. Many abiotic forces influence where life can exist and the types of organisms found in different parts of the biosphere. The abiotic factors influence the distribution of biomes: large areas of land with similar climate, flora, and fauna.
Biogeography
Biogeography is the study of the geographic distribution of living things and the abiotic factors that affect their distribution. Abiotic factors such as temperature and rainfall vary based mainly on latitude and elevation. As these abiotic factors change, the composition of plant and animal communities also changes. For example, if you were to begin a journey at the equator and walk north, you would notice gradual changes in plant communities. At the beginning of your journey, you would see tropical wet forests with broad-leaved evergreen trees, which are characteristic of plant communities found near the equator. As you continued to travel north, you would see these broad-leaved evergreen plants eventually give rise to seasonally dry forests with scattered trees. You would also begin to notice changes in temperature and moisture. At about 30 degrees north, these forests would give way to deserts, which are characterized by low precipitation.
Moving farther north, you would see that deserts are replaced by grasslands or prairies. Eventually, grasslands are replaced by deciduous temperate forests. These deciduous forests give way to the boreal forests found in the subarctic, the area south of the Arctic Circle. Finally, you would reach the Arctic tundra, which is found at the most northern latitudes. This trek north reveals gradual changes in both climate and the types of organisms that have adapted to environmental factors associated with ecosystems found at different latitudes. However, different ecosystems exist at the same latitude due in part to abiotic factors such as jet streams, the Gulf Stream, and ocean currents. If you were to hike up a mountain, the changes you would see in the vegetation would parallel those as you move to higher latitudes.
Ecologists who study biogeography examine patterns of species distribution. No species exists everywhere; for example, the Venus flytrap is endemic to a small area in North and South Carolina. An endemic species is one which is naturally found only in a specific geographic area that is usually restricted in size. Other species are generalists: species which live in a wide variety of geographic areas; the raccoon, for example, is native to most of North and Central America.
Species distribution patterns are based on biotic and abiotic factors and their influences during the very long periods of time required for species evolution; therefore, early studies of biogeography were closely linked to the emergence of evolutionary thinking in the eighteenth century. Some of the most distinctive assemblages of plants and animals occur in regions that have been physically separated for millions of years by geographic barriers. Biologists estimate that Australia, for example, has between 600,000 and 700,000 species of plants and animals. Approximately 3/4 of living plant and mammal species are endemic species found solely in Australia (Figure \(1\)).
Sometimes ecologists discover unique patterns of species distribution by determining where species are not found. Hawaii, for example, has no native land species of reptiles or amphibians, and has only one native terrestrial mammal, the hoary bat. Most of New Guinea, as another example, lacks placental mammals.
Link to Learning
Check out this video to observe a platypus swimming in its natural habitat in New South Wales, Australia.
Plants can be endemic or generalists: endemic plants are found only on specific regions of the Earth, while generalists are found on many regions. Isolated land masses—such as Australia, Hawaii, and Madagascar—often have large numbers of endemic plant species. Some of these plants are endangered due to human activity. The forest gardenia (Gardenia brighamii), for instance, is endemic to Hawaii; only an estimated 15–20 trees are thought to exist (Figure \(2\)).
Energy Sources
Energy from the sun is captured by green plants, algae, cyanobacteria, and photosynthetic protists. These organisms convert solar energy into the chemical energy needed by all living things. Light availability can be an important force directly affecting the evolution of adaptations in photosynthesizers. For instance, plants in the understory of a temperate forest are shaded when the trees above them in the canopy completely leaf out in the late spring. Not surprisingly, understory plants have adaptations to successfully capture available light. One such adaptation is the rapid growth of spring ephemeral plants such as the spring beauty (Figure \(3\)). These spring flowers achieve much of their growth and finish their life cycle (reproduce) early in the season before the trees in the canopy develop leaves.
In aquatic ecosystems, the availability of light may be limited because sunlight is absorbed by water, plants, suspended particles, and resident microorganisms. Toward the bottom of a lake, pond, or ocean, there is a zone that light cannot reach. Photosynthesis cannot take place there and, as a result, a number of adaptations have evolved that enable living things to survive without light. For instance, aquatic plants have photosynthetic tissue near the surface of the water; for example, think of the broad, floating leaves of a water lily—water lilies cannot survive without light. In environments such as hydrothermal vents, some bacteria extract energy from inorganic chemicals because there is no light for photosynthesis.
The availability of nutrients in aquatic systems is also an important aspect of energy or photosynthesis. Many organisms sink to the bottom of the ocean when they die in the open water; when this occurs, the energy found in that living organism is sequestered for some time unless ocean upwelling occurs. Ocean upwelling is the rising of deep ocean waters that occurs when prevailing winds blow along surface waters near a coastline (Figure \(4\)). As the wind pushes ocean waters offshore, water from the bottom of the ocean moves up to replace this water. As a result, the nutrients once contained in dead organisms become available for reuse by other living organisms.
In freshwater systems, the recycling of nutrients occurs in response to air temperature changes. The nutrients at the bottom of lakes are recycled twice each year: in the spring and fall turnover. The spring and fall turnover is a seasonal process that recycles nutrients and oxygen from the bottom of a freshwater ecosystem to the top of a body of water (Figure \(5\)). These turnovers are caused by the formation of a thermocline: a layer of water with a temperature that is significantly different from that of the surrounding layers. In wintertime, the surface of lakes found in many northern regions is frozen. However, the water under the ice is slightly warmer, and the water at the bottom of the lake is warmer yet at 4 °C to 5 °C (39.2 °F to 41 °F). Water is densest at 4 °C; therefore, the deepest water is also the densest. The deepest water is oxygen poor because the decomposition of organic material at the bottom of the lake uses up available oxygen that cannot be replaced by means of oxygen diffusion into the water due to the surface ice layer.
Art Connection
How might turnover in tropical lakes differ from turnover in lakes that exist in temperate regions?
In springtime, air temperatures increase and surface ice melts. When the temperature of the surface water begins to reach 4 °C, the water becomes heavier and sinks to the bottom. The water at the bottom of the lake is then displaced by the heavier surface water and, thus, rises to the top. As that water rises to the top, the sediments and nutrients from the lake bottom are brought along with it. During the summer months, the lake water stratifies, or forms layers, with the warmest water at the lake surface.
As air temperatures drop in the fall, the temperature of the lake water cools to 4 °C; therefore, this causes fall turnover as the heavy cold water sinks and displaces the water at the bottom. The oxygen-rich water at the surface of the lake then moves to the bottom of the lake, while the nutrients at the bottom of the lake rise to the surface (Figure \(5\)). During the winter, the oxygen at the bottom of the lake is used by decomposers and other organisms requiring oxygen, such as fish.
Temperature
Temperature affects the physiology of living things as well as the density and state of water. Temperature exerts an important influence on living things because few living things can survive at temperatures below 0 °C (32 °F) due to metabolic constraints. It is also rare for living things to survive at temperatures exceeding 45 °C (113 °F); this is a reflection of evolutionary response to typical temperatures. Enzymes are most efficient within a narrow and specific range of temperatures; enzyme degradation can occur at higher temperatures. Therefore, organisms either must maintain an internal temperature or they must inhabit an environment that will keep the body within a temperature range that supports metabolism. Some animals have adapted to enable their bodies to survive significant temperature fluctuations, such as seen in hibernation or reptilian torpor. Similarly, some bacteria are adapted to surviving in extremely hot temperatures such as geysers. Such bacteria are examples of extremophiles: organisms that thrive in extreme environments.
Temperature can limit the distribution of living things. Animals faced with temperature fluctuations may respond with adaptations, such as migration, in order to survive. Migration, the movement from one place to another, is an adaptation found in many animals, including many that inhabit seasonally cold climates. Migration solves problems related to temperature, locating food, and finding a mate. In migration, for instance, the Arctic Tern (Sterna paradisaea) makes a 40,000 km (24,000 mi) round trip flight each year between its feeding grounds in the southern hemisphere and its breeding grounds in the Arctic Ocean. Monarch butterflies (Danaus plexippus) live in the eastern United States in the warmer months and migrate to Mexico and the southern United States in the wintertime. Some species of mammals also make migratory forays. Reindeer (Rangifer tarandus) travel about 5,000 km (3,100 mi) each year to find food. Amphibians and reptiles are more limited in their distribution because they lack migratory ability. Not all animals that can migrate do so: migration carries risk and comes at a high energy cost.
Some animals hibernate or estivate to survive hostile temperatures. Hibernation enables animals to survive cold conditions, and estivation allows animals to survive the hostile conditions of a hot, dry climate. Animals that hibernate or estivate enter a state known as torpor: a condition in which their metabolic rate is significantly lowered. This enables the animal to wait until its environment better supports its survival. Some amphibians, such as the wood frog (Rana sylvatica), have an antifreeze-like chemical in their cells, which retains the cells’ integrity and prevents them from bursting.
Water
Water is required by all living things because it is critical for cellular processes. Since terrestrial organisms lose water to the environment by simple diffusion, they have evolved many adaptations to retain water.
• Plants have a number of interesting features on their leaves, such as leaf hairs and a waxy cuticle, that serve to decrease the rate of water loss via transpiration.
• Freshwater organisms are surrounded by water and are constantly in danger of having water rush into their cells because of osmosis. Many adaptations of organisms living in freshwater environments have evolved to ensure that solute concentrations in their bodies remain within appropriate levels. One such adaptation is the excretion of dilute urine.
• Marine organisms are surrounded by water with a higher solute concentration than the organism and, thus, are in danger of losing water to the environment because of osmosis. These organisms have morphological and physiological adaptations to retain water and release solutes into the environment. For example, Marine iguanas (Amblyrhynchus cristatus), sneeze out water vapor that is high in salt in order to maintain solute concentrations within an acceptable range while swimming in the ocean and eating marine plants.
Inorganic Nutrients and Soil
Inorganic nutrients, such as nitrogen and phosphorus, are important in the distribution and the abundance of living things. Plants obtain these inorganic nutrients from the soil when water moves into the plant through the roots. Therefore, soil structure (particle size of soil components), soil pH, and soil nutrient content play an important role in the distribution of plants. Animals obtain inorganic nutrients from the food they consume. Therefore, animal distributions are related to the distribution of what they eat. In some cases, animals will follow their food resource as it moves through the environment.
Other Aquatic Factors
Some abiotic factors, such as oxygen, are important in aquatic ecosystems as well as terrestrial environments. Terrestrial animals obtain oxygen from the air they breathe. Oxygen availability can be an issue for organisms living at very high elevations, however, where there are fewer molecules of oxygen in the air. In aquatic systems, the concentration of dissolved oxygen is related to water temperature and the speed at which the water moves. Cold water has more dissolved oxygen than warmer water. In addition, salinity, current, and tide can be important abiotic factors in aquatic ecosystems.
Other Terrestrial Factors
Wind can be an important abiotic factor because it influences the rate of evaporation and transpiration. The physical force of wind is also important because it can move soil, water, or other abiotic factors, as well as an ecosystem’s organisms.
Fire is another terrestrial factor that can be an important agent of disturbance in terrestrial ecosystems. Some organisms are adapted to fire and, thus, require the high heat associated with fire to complete a part of their life cycle. For example, the jack pine—a coniferous tree—requires heat from fire for its seed cones to open (Figure \(6\)). Through the burning of pine needles, fire adds nitrogen to the soil and limits competition by destroying undergrowth.
Abiotic Factors Influencing Plant Growth
Temperature and moisture are important influences on plant production (primary productivity) and the amount of organic matter available as food (net primary productivity). Net primary productivity is an estimation of all of the organic matter available as food; it is calculated as the total amount of carbon fixed per year minus the amount that is oxidized during cellular respiration. In terrestrial environments, net primary productivity is estimated by measuring the aboveground biomass per unit area, which is the total mass of living plants, excluding roots. This means that a large percentage of plant biomass which exists underground is not included in this measurement. Net primary productivity is an important variable when considering differences in biomes. Very productive biomes have a high level of aboveground biomass.
Annual biomass production is directly related to the abiotic components of the environment. Environments with the greatest amount of biomass have conditions in which photosynthesis, plant growth, and the resulting net primary productivity are optimized. The climate of these areas is warm and wet. Photosynthesis can proceed at a high rate, enzymes can work most efficiently, and stomata can remain open without the risk of excessive transpiration; together, these factors lead to the maximal amount of carbon dioxide (CO2) moving into the plant, resulting in high biomass production. The aboveground biomass produces several important resources for other living things, including habitat and food. Conversely, dry and cold environments have lower photosynthetic rates and therefore less biomass. The animal communities living there will also be affected by the decrease in available food.
Summary
Biogeography is the study of the geographic distribution of living things and the abiotic factors that affect their distribution. Endemic species are species that are naturally found only in a specific geographic area. The distribution of living things is influenced by several environmental factors that are, in part, controlled by the latitude or elevation at which an organism is found. Ocean upwelling and spring and fall turnovers are important processes regulating the distribution of nutrients and other abiotic factors important in aquatic ecosystems. Energy sources, temperature, water, inorganic nutrients, and soil are factors limiting the distribution of living things in terrestrial systems. Net primary productivity is a measure of the amount of biomass produced by a biome.
Art Connections
Figure \(5\): How might turnover in tropical lakes differ from turnover in lakes that exist in temperate regions?
Answer
Tropical lakes don’t freeze, so they don’t undergo spring turnover in the same way temperate lakes do. However, stratification does occur, as well as seasonal turnover.
Glossary
aboveground biomass
total mass of aboveground living plants per area
biogeography
study of the geographic distribution of living things and the abiotic factors that affect their distribution
biome
ecological community of plants, animals, and other organisms that is adapted to a characteristic set of environmental conditions
endemic
species found only in a specific geographic area that is usually restricted in size
fall and spring turnover
seasonal process that recycles nutrients and oxygen from the bottom of a freshwater ecosystem to the top
net primary productivity
measurement of the energy accumulation within an ecosystem, calculated as the total amount of carbon fixed per year minus the amount that is oxidized during cellular respiration
ocean upwelling
rising of deep ocean waters that occurs when prevailing winds blow along surface waters near a coastline
thermocline
layer of water with a temperature that is significantly different from that of the surrounding layers | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/55%3A_Community_Ecology/55.01%3A_Biological_Communities-_Species_Living_Together/55.1.01%3A_Biogeography.txt |
Skills to Develop
• Discuss the predator-prey cycle
• Give examples of defenses against predation and herbivory
• Describe the competitive exclusion principle
• Give examples of symbiotic relationships between species
• Describe community structure and succession
Populations rarely, if ever, live in isolation from populations of other species. In most cases, numerous species share a habitat. The interactions between these populations play a major role in regulating population growth and abundance. All populations occupying the same habitat form a community: populations inhabiting a specific area at the same time. The number of species occupying the same habitat and their relative abundance is known as species diversity. Areas with low diversity, such as the glaciers of Antarctica, still contain a wide variety of living things, whereas the diversity of tropical rainforests is so great that it cannot be counted. Ecology is studied at the community level to understand how species interact with each other and compete for the same resources.
Predation and Herbivory
Perhaps the classical example of species interaction is predation: the hunting of prey by its predator. Nature shows on television highlight the drama of one living organism killing another. Populations of predators and prey in a community are not constant over time: in most cases, they vary in cycles that appear to be related. The most often cited example of predator-prey dynamics is seen in the cycling of the lynx (predator) and the snowshoe hare (prey), using nearly 200 year-old trapping data from North American forests (Figure \(1\)). This cycle of predator and prey lasts approximately 10 years, with the predator population lagging 1–2 years behind that of the prey population. As the hare numbers increase, there is more food available for the lynx, allowing the lynx population to increase as well. When the lynx population grows to a threshold level, however, they kill so many hares that hare population begins to decline, followed by a decline in the lynx population because of scarcity of food. When the lynx population is low, the hare population size begins to increase due, at least in part, to low predation pressure, starting the cycle anew.
The idea that the population cycling of the two species is entirely controlled by predation models has come under question. More recent studies have pointed to undefined density-dependent factors as being important in the cycling, in addition to predation. One possibility is that the cycling is inherent in the hare population due to density-dependent effects such as lower fecundity (maternal stress) caused by crowding when the hare population gets too dense. The hare cycling would then induce the cycling of the lynx because it is the lynxes’ major food source. The more we study communities, the more complexities we find, allowing ecologists to derive more accurate and sophisticated models of population dynamics.
Herbivory describes the consumption of plants by insects and other animals, and it is another interspecific relationship that affects populations. Unlike animals, most plants cannot outrun predators or use mimicry to hide from hungry animals. Some plants have developed mechanisms to defend against herbivory. Other species have developed mutualistic relationships; for example, herbivory provides a mechanism of seed distribution that aids in plant reproduction.
Defense Mechanisms against Predation and Herbivory
The study of communities must consider evolutionary forces that act on the members of the various populations contained within it. Species are not static, but slowly changing and adapting to their environment by natural selection and other evolutionary forces. Species have evolved numerous mechanisms to escape predation and herbivory. These defenses may be mechanical, chemical, physical, or behavioral.
Mechanical defenses, such as the presence of thorns on plants or the hard shell on turtles, discourage animal predation and herbivory by causing physical pain to the predator or by physically preventing the predator from being able to eat the prey. Chemical defenses are produced by many animals as well as plants, such as the foxglove which is extremely toxic when eaten. Figure \(2\) shows some organisms’ defenses against predation and herbivory.
Many species use their body shape and coloration to avoid being detected by predators. The tropical walking stick is an insect with the coloration and body shape of a twig which makes it very hard to see when stationary against a background of real twigs (Figure \(3\)a). In another example, the chameleon can change its color to match its surroundings (Figure \(3\)b). Both of these are examples of camouflage, or avoiding detection by blending in with the background.
Some species use coloration as a way of warning predators that they are not good to eat. For example, the cinnabar moth caterpillar, the fire-bellied toad, and many species of beetle have bright colors that warn of a foul taste, the presence of toxic chemical, and/or the ability to sting or bite, respectively. Predators that ignore this coloration and eat the organisms will experience their unpleasant taste or presence of toxic chemicals and learn not to eat them in the future. This type of defensive mechanism is called aposematic coloration, or warning coloration (Figure \(4\)).
While some predators learn to avoid eating certain potential prey because of their coloration, other species have evolved mechanisms to mimic this coloration to avoid being eaten, even though they themselves may not be unpleasant to eat or contain toxic chemicals. In Batesian mimicry, a harmless species imitates the warning coloration of a harmful one. Assuming they share the same predators, this coloration then protects the harmless ones, even though they do not have the same level of physical or chemical defenses against predation as the organism they mimic. Many insect species mimic the coloration of wasps or bees, which are stinging, venomous insects, thereby discouraging predation (Figure \(5\)).
In Müllerian mimicry, multiple species share the same warning coloration, but all of them actually have defenses. Figure \(6\) shows a variety of foul-tasting butterflies with similar coloration. In Emsleyan/Mertensian mimicry, a deadly prey mimics a less dangerous one, such as the venomous coral snake mimicking the non-venomous milk snake. This type of mimicry is extremely rare and more difficult to understand than the previous two types. For this type of mimicry to work, it is essential that eating the milk snake has unpleasant but not fatal consequences. Then, these predators learn not to eat snakes with this coloration, protecting the coral snake as well. If the snake were fatal to the predator, there would be no opportunity for the predator to learn not to eat it, and the benefit for the less toxic species would disappear.
Link to Learning
Go to this website to view stunning examples of mimicry.
Competitive Exclusion Principle
Resources are often limited within a habitat and multiple species may compete to obtain them. All species have an ecological niche in the ecosystem, which describes how they acquire the resources they need and how they interact with other species in the community. The competitive exclusion principle states that two species cannot occupy the same niche in a habitat. In other words, different species cannot coexist in a community if they are competing for all the same resources. An example of this principle is shown in Figure \(7\), with two protozoan species, Paramecium aurelia and Paramecium caudatum. When grown individually in the laboratory, they both thrive. But when they are placed together in the same test tube (habitat), P. aurelia outcompetes P. caudatum for food, leading to the latter’s eventual extinction.
This exclusion may be avoided if a population evolves to make use of a different resource, a different area of the habitat, or feeds during a different time of day, called resource partitioning. The two organisms are then said to occupy different microniches. These organisms coexist by minimizing direct competition.
Symbiosis
Symbiotic relationships, or symbioses (plural), are close interactions between individuals of different species over an extended period of time which impact the abundance and distribution of the associating populations. Most scientists accept this definition, but some restrict the term to only those species that are mutualistic, where both individuals benefit from the interaction. In this discussion, the broader definition will be used.
Commensalism
A commensal relationship occurs when one species benefits from the close, prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure \(8\)). The tree is not harmed by the presence of the nest among its branches. The nests are light and produce little strain on the structural integrity of the branch, and most of the leaves, which the tree uses to get energy by photosynthesis, are above the nest so they are unaffected. The bird, on the other hand, benefits greatly. If the bird had to nest in the open, its eggs and young would be vulnerable to predators. Another example of a commensal relationship is the clown fish and the sea anemone. The sea anemone is not harmed by the fish, and the fish benefits with protection from predators who would be stung upon nearing the sea anemone.
Mutualism
A second type of symbiotic relationship is called mutualism, where two species benefit from their interaction. Some scientists believe that these are the only true examples of symbiosis. For example, termites have a mutualistic relationship with protozoa that live in the insect’s gut (Figure \(9\)a). The termite benefits from the ability of bacterial symbionts within the protozoa to digest cellulose. The termite itself cannot do this, and without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa and the bacterial symbionts benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. Lichens have a mutualistic relationship between fungus and photosynthetic algae or bacteria (Figure \(9\)b). As these symbionts grow together, the glucose produced by the algae provides nourishment for both organisms, whereas the physical structure of the lichen protects the algae from the elements and makes certain nutrients in the atmosphere more available to the algae.
Parasitism
A parasite is an organism that lives in or on another living organism and derives nutrients from it. In this relationship, the parasite benefits, but the organism being fed upon, the host, is harmed. The host is usually weakened by the parasite as it siphons resources the host would normally use to maintain itself. The parasite, however, is unlikely to kill the host, especially not quickly, because this would allow no time for the organism to complete its reproductive cycle by spreading to another host.
The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm is a parasite that causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure \(10\)). The tapeworm can live inside the intestine of the host for several years, benefiting from the food the host is bringing into its gut by eating, and may grow to be over 50 ft long by adding segments. The parasite moves from species to species in a cycle, making two hosts necessary to complete its life cycle. Another common parasite is Plasmodium falciparum, the protozoan cause of malaria, a significant disease in many parts of the world. Living in human liver and red blood cells, the organism reproduces asexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to human by mosquitoes, one of many arthropod-borne infectious diseases.
Characteristics of Communities
Communities are complex entities that can be characterized by their structure (the types and numbers of species present) and dynamics (how communities change over time). Understanding community structure and dynamics enables community ecologists to manage ecosystems more effectively.
Foundation Species
Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are usually the primary producers: organisms that bring most of the energy into the community. Kelp, brown algae, is a foundation species, forming the basis of the kelp forests off the coast of California.
Foundation species may physically modify the environment to produce and maintain habitats that benefit the other organisms that use them. An example is the photosynthetic corals of the coral reef (Figure \(11\)). Corals themselves are not photosynthetic, but harbor symbionts within their body tissues (dinoflagellates called zooxanthellae) that perform photosynthesis; this is another example of a mutualism. The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.
Biodiversity, Species Richness, and Relative Species Abundance
Biodiversity describes a community’s biological complexity: it is measured by the number of different species (species richness) in a particular area and their relative abundance (species evenness). The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term that is used to describe the number of species living in a habitat or biome. Species richness varies across the globe (Figure \(12\)). One factor in determining species richness is latitude, with the greatest species richness occurring in ecosystems near the equator, which often have warmer temperatures, large amounts of rainfall, and low seasonality. The lowest species richness occurs near the poles, which are much colder, drier, and thus less conducive to life in Geologic time (time since glaciations). The predictability of climate or productivity is also an important factor. Other factors influence species richness as well. For example, the study of island biogeography attempts to explain the relatively high species richness found in certain isolated island chains, including the Galápagos Islands that inspired the young Darwin. Relative species abundance is the number of individuals in a species relative to the total number of individuals in all species within a habitat, ecosystem, or biome. Foundation species often have the highest relative abundance of species.
Keystone Species
A keystone species is one whose presence is key to maintaining biodiversity within an ecosystem and to upholding an ecological community’s structure. The intertidal sea star, Pisaster ochraceus, of the northwestern United States is a keystone species (Figure \(13\)). Studies have shown that when this organism is removed from communities, populations of their natural prey (mussels) increase, completely altering the species composition and reducing biodiversity. Another keystone species is the banded tetra, a fish in tropical streams, which supplies nearly all of the phosphorus, a necessary inorganic nutrient, to the rest of the community. If these fish were to become extinct, the community would be greatly affected.
Everyday Connection: Invasive Species
Invasive species are non-native organisms that, when introduced to an area out of their native range, threaten the ecosystem balance of that habitat. Many such species exist in the United States, as shown in Figure \(14\). Whether enjoying a forest hike, taking a summer boat trip, or simply walking down an urban street, you have likely encountered an invasive species.
One of the many recent proliferations of an invasive species concerns the growth of Asian carp populations. Asian carp were introduced to the United States in the 1970s by fisheries and sewage treatment facilities that used the fish’s excellent filter feeding capabilities to clean their ponds of excess plankton. Some of the fish escaped, however, and by the 1980s they had colonized many waterways of the Mississippi River basin, including the Illinois and Missouri Rivers.
Voracious eaters and rapid reproducers, Asian carp may outcompete native species for food, potentially leading to their extinction. For example, black carp are voracious eaters of native mussels and snails, limiting this food source for native fish species. Silver carp eat plankton that native mussels and snails feed on, reducing this food source by a different alteration of the food web. In some areas of the Mississippi River, Asian carp species have become the most predominant, effectively outcompeting native fishes for habitat. In some parts of the Illinois River, Asian carp constitute 95 percent of the community's biomass. Although edible, the fish is bony and not a desired food in the United States. Moreover, their presence threatens the native fish and fisheries of the Great Lakes, which are important to local economies and recreational anglers. Asian carp have even injured humans. The fish, frightened by the sound of approaching motorboats, thrust themselves into the air, often landing in the boat or directly hitting the boaters.
The Great Lakes and their prized salmon and lake trout fisheries are also being threatened by these invasive fish. Asian carp have already colonized rivers and canals that lead into Lake Michigan. One infested waterway of particular importance is the Chicago Sanitary and Ship Channel, the major supply waterway linking the Great Lakes to the Mississippi River. To prevent the Asian carp from leaving the canal, a series of electric barriers have been successfully used to discourage their migration; however, the threat is significant enough that several states and Canada have sued to have the Chicago channel permanently cut off from Lake Michigan. Local and national politicians have weighed in on how to solve the problem, but no one knows whether the Asian carp will ultimately be considered a nuisance, like other invasive species such as the water hyacinth and zebra mussel, or whether it will be the destroyer of the largest freshwater fishery of the world.
The issues associated with Asian carp show how population and community ecology, fisheries management, and politics intersect on issues of vital importance to the human food supply and economy. Socio-political issues like this make extensive use of the sciences of population ecology (the study of members of a particular species occupying a particular area known as a habitat) and community ecology (the study of the interaction of all species within a habitat).
Community Dynamics
Community dynamics are the changes in community structure and composition over time. Sometimes these changes are induced by environmental disturbances such as volcanoes, earthquakes, storms, fires, and climate change. Communities with a stable structure are said to be at equilibrium. Following a disturbance, the community may or may not return to the equilibrium state.
Succession describes the sequential appearance and disappearance of species in a community over time. In primary succession, newly exposed or newly formed land is colonized by living things; in secondary succession, part of an ecosystem is disturbed and remnants of the previous community remain.
Primary Succession and Pioneer Species
Primary succession occurs when new land is formed or rock is exposed: for example, following the eruption of volcanoes, such as those on the Big Island of Hawaii. As lava flows into the ocean, new land is continually being formed. On the Big Island, approximately 32 acres of land is added each year. First, weathering and other natural forces break down the substrate enough for the establishment of certain hearty plants and lichens with few soil requirements, known as pioneer species (Figure \(15\)). These species help to further break down the mineral rich lava into soil where other, less hardy species will grow and eventually replace the pioneer species. In addition, as these early species grow and die, they add to an ever-growing layer of decomposing organic material and contribute to soil formation. Over time the area will reach an equilibrium state, with a set of organisms quite different from the pioneer species.
Secondary succession
A classic example of secondary succession occurs in oak and hickory forests cleared by wildfire (Figure \(16\)). Wildfires will burn most vegetation and kill those animals unable to flee the area. Their nutrients, however, are returned to the ground in the form of ash. Thus, even when areas are devoid of life due to severe fires, the area will soon be ready for new life to take hold.
Before the fire, the vegetation was dominated by tall trees with access to the major plant energy resource: sunlight. Their height gave them access to sunlight while also shading the ground and other low-lying species. After the fire, though, these trees are no longer dominant. Thus, the first plants to grow back are usually annual plants followed within a few years by quickly growing and spreading grasses and other pioneer species. Due to, at least in part, changes in the environment brought on by the growth of the grasses and other species, over many years, shrubs will emerge along with small pine, oak, and hickory trees. These organisms are called intermediate species. Eventually, over 150 years, the forest will reach its equilibrium point where species composition is no longer changing and resembles the community before the fire. This equilibrium state is referred to as the climax community, which will remain stable until the next disturbance.
Summary
Communities include all the different species living in a given area. The variety of these species is called species richness. Many organisms have developed defenses against predation and herbivory, including mechanical defenses, warning coloration, and mimicry, as a result of evolution and the interaction with other members of the community. Two species cannot exist in the same habitat competing directly for the same resources. Species may form symbiotic relationships such as commensalism or mutualism. Community structure is described by its foundation and keystone species. Communities respond to environmental disturbances by succession (the predictable appearance of different types of plant species) until a stable community structure is established.
Glossary
aposematic coloration
warning coloration used as a defensive mechanism against predation
Batesian mimicry
type of mimicry where a non-harmful species takes on the warning colorations of a harmful one
camouflage
avoid detection by blending in with the background.
climax community
final stage of succession, where a stable community is formed by a characteristic assortment of plant and animal species
commensalism
relationship between species wherein one species benefits from the close, prolonged interaction, while the other species neither benefits nor is harmed
competitive exclusion principle
no two species within a habitat can coexist when they compete for the same resources at the same place and time
Emsleyan/Mertensian mimicry
type of mimicry where a harmful species resembles a less harmful one
environmental disturbance
change in the environment caused by natural disasters or human activities
foundation species
species which often forms the major structural portion of the habitat
host
organism a parasite lives on
island biogeography
study of life on island chains and how their geography interacts with the diversity of species found there
keystone species
species whose presence is key to maintaining biodiversity in an ecosystem and to upholding an ecological community’s structure
Müllerian mimicry
type of mimicry where species share warning coloration and all are harmful to predators
mutualism
symbiotic relationship between two species where both species benefit
parasite
organism that uses resources from another species, the host
pioneer species
first species to appear in primary and secondary succession
primary succession
succession on land that previously has had no life
relative species abundance
absolute population size of a particular species relative to the population sizes of other species within the community
secondary succession
succession in response to environmental disturbances that move a community away from its equilibrium
species richness
number of different species in a community
symbiosis
close interaction between individuals of different species over an extended period of time that impacts the abundance and distribution of the associating populations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/55%3A_Community_Ecology/55.01%3A_Biological_Communities-_Species_Living_Together/55.1.02%3A_Community_Ecology.txt |
Skills to Develop
• Discuss the predator-prey cycle
• Give examples of defenses against predation and herbivory
• Describe the competitive exclusion principle
• Give examples of symbiotic relationships between species
• Describe community structure and succession
Populations rarely, if ever, live in isolation from populations of other species. In most cases, numerous species share a habitat. The interactions between these populations play a major role in regulating population growth and abundance. All populations occupying the same habitat form a community: populations inhabiting a specific area at the same time. The number of species occupying the same habitat and their relative abundance is known as species diversity. Areas with low diversity, such as the glaciers of Antarctica, still contain a wide variety of living things, whereas the diversity of tropical rainforests is so great that it cannot be counted. Ecology is studied at the community level to understand how species interact with each other and compete for the same resources.
Predation and Herbivory
Perhaps the classical example of species interaction is predation: the hunting of prey by its predator. Nature shows on television highlight the drama of one living organism killing another. Populations of predators and prey in a community are not constant over time: in most cases, they vary in cycles that appear to be related. The most often cited example of predator-prey dynamics is seen in the cycling of the lynx (predator) and the snowshoe hare (prey), using nearly 200 year-old trapping data from North American forests (Figure \(1\)). This cycle of predator and prey lasts approximately 10 years, with the predator population lagging 1–2 years behind that of the prey population. As the hare numbers increase, there is more food available for the lynx, allowing the lynx population to increase as well. When the lynx population grows to a threshold level, however, they kill so many hares that hare population begins to decline, followed by a decline in the lynx population because of scarcity of food. When the lynx population is low, the hare population size begins to increase due, at least in part, to low predation pressure, starting the cycle anew.
The idea that the population cycling of the two species is entirely controlled by predation models has come under question. More recent studies have pointed to undefined density-dependent factors as being important in the cycling, in addition to predation. One possibility is that the cycling is inherent in the hare population due to density-dependent effects such as lower fecundity (maternal stress) caused by crowding when the hare population gets too dense. The hare cycling would then induce the cycling of the lynx because it is the lynxes’ major food source. The more we study communities, the more complexities we find, allowing ecologists to derive more accurate and sophisticated models of population dynamics.
Herbivory describes the consumption of plants by insects and other animals, and it is another interspecific relationship that affects populations. Unlike animals, most plants cannot outrun predators or use mimicry to hide from hungry animals. Some plants have developed mechanisms to defend against herbivory. Other species have developed mutualistic relationships; for example, herbivory provides a mechanism of seed distribution that aids in plant reproduction.
Defense Mechanisms against Predation and Herbivory
The study of communities must consider evolutionary forces that act on the members of the various populations contained within it. Species are not static, but slowly changing and adapting to their environment by natural selection and other evolutionary forces. Species have evolved numerous mechanisms to escape predation and herbivory. These defenses may be mechanical, chemical, physical, or behavioral.
Mechanical defenses, such as the presence of thorns on plants or the hard shell on turtles, discourage animal predation and herbivory by causing physical pain to the predator or by physically preventing the predator from being able to eat the prey. Chemical defenses are produced by many animals as well as plants, such as the foxglove which is extremely toxic when eaten. Figure \(2\) shows some organisms’ defenses against predation and herbivory.
Many species use their body shape and coloration to avoid being detected by predators. The tropical walking stick is an insect with the coloration and body shape of a twig which makes it very hard to see when stationary against a background of real twigs (Figure \(3\)a). In another example, the chameleon can change its color to match its surroundings (Figure \(3\)b). Both of these are examples of camouflage, or avoiding detection by blending in with the background.
Some species use coloration as a way of warning predators that they are not good to eat. For example, the cinnabar moth caterpillar, the fire-bellied toad, and many species of beetle have bright colors that warn of a foul taste, the presence of toxic chemical, and/or the ability to sting or bite, respectively. Predators that ignore this coloration and eat the organisms will experience their unpleasant taste or presence of toxic chemicals and learn not to eat them in the future. This type of defensive mechanism is called aposematic coloration, or warning coloration (Figure \(4\)).
While some predators learn to avoid eating certain potential prey because of their coloration, other species have evolved mechanisms to mimic this coloration to avoid being eaten, even though they themselves may not be unpleasant to eat or contain toxic chemicals. In Batesian mimicry, a harmless species imitates the warning coloration of a harmful one. Assuming they share the same predators, this coloration then protects the harmless ones, even though they do not have the same level of physical or chemical defenses against predation as the organism they mimic. Many insect species mimic the coloration of wasps or bees, which are stinging, venomous insects, thereby discouraging predation (Figure \(5\)).
In Müllerian mimicry, multiple species share the same warning coloration, but all of them actually have defenses. Figure \(6\) shows a variety of foul-tasting butterflies with similar coloration. In Emsleyan/Mertensian mimicry, a deadly prey mimics a less dangerous one, such as the venomous coral snake mimicking the non-venomous milk snake. This type of mimicry is extremely rare and more difficult to understand than the previous two types. For this type of mimicry to work, it is essential that eating the milk snake has unpleasant but not fatal consequences. Then, these predators learn not to eat snakes with this coloration, protecting the coral snake as well. If the snake were fatal to the predator, there would be no opportunity for the predator to learn not to eat it, and the benefit for the less toxic species would disappear.
Link to Learning
Go to this website to view stunning examples of mimicry.
Competitive Exclusion Principle
Resources are often limited within a habitat and multiple species may compete to obtain them. All species have an ecological niche in the ecosystem, which describes how they acquire the resources they need and how they interact with other species in the community. The competitive exclusion principle states that two species cannot occupy the same niche in a habitat. In other words, different species cannot coexist in a community if they are competing for all the same resources. An example of this principle is shown in Figure \(7\), with two protozoan species, Paramecium aurelia and Paramecium caudatum. When grown individually in the laboratory, they both thrive. But when they are placed together in the same test tube (habitat), P. aurelia outcompetes P. caudatum for food, leading to the latter’s eventual extinction.
This exclusion may be avoided if a population evolves to make use of a different resource, a different area of the habitat, or feeds during a different time of day, called resource partitioning. The two organisms are then said to occupy different microniches. These organisms coexist by minimizing direct competition.
Symbiosis
Symbiotic relationships, or symbioses (plural), are close interactions between individuals of different species over an extended period of time which impact the abundance and distribution of the associating populations. Most scientists accept this definition, but some restrict the term to only those species that are mutualistic, where both individuals benefit from the interaction. In this discussion, the broader definition will be used.
Commensalism
A commensal relationship occurs when one species benefits from the close, prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure \(8\)). The tree is not harmed by the presence of the nest among its branches. The nests are light and produce little strain on the structural integrity of the branch, and most of the leaves, which the tree uses to get energy by photosynthesis, are above the nest so they are unaffected. The bird, on the other hand, benefits greatly. If the bird had to nest in the open, its eggs and young would be vulnerable to predators. Another example of a commensal relationship is the clown fish and the sea anemone. The sea anemone is not harmed by the fish, and the fish benefits with protection from predators who would be stung upon nearing the sea anemone.
Mutualism
A second type of symbiotic relationship is called mutualism, where two species benefit from their interaction. Some scientists believe that these are the only true examples of symbiosis. For example, termites have a mutualistic relationship with protozoa that live in the insect’s gut (Figure \(9\)a). The termite benefits from the ability of bacterial symbionts within the protozoa to digest cellulose. The termite itself cannot do this, and without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa and the bacterial symbionts benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. Lichens have a mutualistic relationship between fungus and photosynthetic algae or bacteria (Figure \(9\)b). As these symbionts grow together, the glucose produced by the algae provides nourishment for both organisms, whereas the physical structure of the lichen protects the algae from the elements and makes certain nutrients in the atmosphere more available to the algae.
Parasitism
A parasite is an organism that lives in or on another living organism and derives nutrients from it. In this relationship, the parasite benefits, but the organism being fed upon, the host, is harmed. The host is usually weakened by the parasite as it siphons resources the host would normally use to maintain itself. The parasite, however, is unlikely to kill the host, especially not quickly, because this would allow no time for the organism to complete its reproductive cycle by spreading to another host.
The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm is a parasite that causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure \(10\)). The tapeworm can live inside the intestine of the host for several years, benefiting from the food the host is bringing into its gut by eating, and may grow to be over 50 ft long by adding segments. The parasite moves from species to species in a cycle, making two hosts necessary to complete its life cycle. Another common parasite is Plasmodium falciparum, the protozoan cause of malaria, a significant disease in many parts of the world. Living in human liver and red blood cells, the organism reproduces asexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to human by mosquitoes, one of many arthropod-borne infectious diseases.
Characteristics of Communities
Communities are complex entities that can be characterized by their structure (the types and numbers of species present) and dynamics (how communities change over time). Understanding community structure and dynamics enables community ecologists to manage ecosystems more effectively.
Foundation Species
Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are usually the primary producers: organisms that bring most of the energy into the community. Kelp, brown algae, is a foundation species, forming the basis of the kelp forests off the coast of California.
Foundation species may physically modify the environment to produce and maintain habitats that benefit the other organisms that use them. An example is the photosynthetic corals of the coral reef (Figure \(11\)). Corals themselves are not photosynthetic, but harbor symbionts within their body tissues (dinoflagellates called zooxanthellae) that perform photosynthesis; this is another example of a mutualism. The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.
Biodiversity, Species Richness, and Relative Species Abundance
Biodiversity describes a community’s biological complexity: it is measured by the number of different species (species richness) in a particular area and their relative abundance (species evenness). The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term that is used to describe the number of species living in a habitat or biome. Species richness varies across the globe (Figure \(12\)). One factor in determining species richness is latitude, with the greatest species richness occurring in ecosystems near the equator, which often have warmer temperatures, large amounts of rainfall, and low seasonality. The lowest species richness occurs near the poles, which are much colder, drier, and thus less conducive to life in Geologic time (time since glaciations). The predictability of climate or productivity is also an important factor. Other factors influence species richness as well. For example, the study of island biogeography attempts to explain the relatively high species richness found in certain isolated island chains, including the Galápagos Islands that inspired the young Darwin. Relative species abundance is the number of individuals in a species relative to the total number of individuals in all species within a habitat, ecosystem, or biome. Foundation species often have the highest relative abundance of species.
Keystone Species
A keystone species is one whose presence is key to maintaining biodiversity within an ecosystem and to upholding an ecological community’s structure. The intertidal sea star, Pisaster ochraceus, of the northwestern United States is a keystone species (Figure \(13\)). Studies have shown that when this organism is removed from communities, populations of their natural prey (mussels) increase, completely altering the species composition and reducing biodiversity. Another keystone species is the banded tetra, a fish in tropical streams, which supplies nearly all of the phosphorus, a necessary inorganic nutrient, to the rest of the community. If these fish were to become extinct, the community would be greatly affected.
Everyday Connection: Invasive Species
Invasive species are non-native organisms that, when introduced to an area out of their native range, threaten the ecosystem balance of that habitat. Many such species exist in the United States, as shown in Figure \(14\). Whether enjoying a forest hike, taking a summer boat trip, or simply walking down an urban street, you have likely encountered an invasive species.
One of the many recent proliferations of an invasive species concerns the growth of Asian carp populations. Asian carp were introduced to the United States in the 1970s by fisheries and sewage treatment facilities that used the fish’s excellent filter feeding capabilities to clean their ponds of excess plankton. Some of the fish escaped, however, and by the 1980s they had colonized many waterways of the Mississippi River basin, including the Illinois and Missouri Rivers.
Voracious eaters and rapid reproducers, Asian carp may outcompete native species for food, potentially leading to their extinction. For example, black carp are voracious eaters of native mussels and snails, limiting this food source for native fish species. Silver carp eat plankton that native mussels and snails feed on, reducing this food source by a different alteration of the food web. In some areas of the Mississippi River, Asian carp species have become the most predominant, effectively outcompeting native fishes for habitat. In some parts of the Illinois River, Asian carp constitute 95 percent of the community's biomass. Although edible, the fish is bony and not a desired food in the United States. Moreover, their presence threatens the native fish and fisheries of the Great Lakes, which are important to local economies and recreational anglers. Asian carp have even injured humans. The fish, frightened by the sound of approaching motorboats, thrust themselves into the air, often landing in the boat or directly hitting the boaters.
The Great Lakes and their prized salmon and lake trout fisheries are also being threatened by these invasive fish. Asian carp have already colonized rivers and canals that lead into Lake Michigan. One infested waterway of particular importance is the Chicago Sanitary and Ship Channel, the major supply waterway linking the Great Lakes to the Mississippi River. To prevent the Asian carp from leaving the canal, a series of electric barriers have been successfully used to discourage their migration; however, the threat is significant enough that several states and Canada have sued to have the Chicago channel permanently cut off from Lake Michigan. Local and national politicians have weighed in on how to solve the problem, but no one knows whether the Asian carp will ultimately be considered a nuisance, like other invasive species such as the water hyacinth and zebra mussel, or whether it will be the destroyer of the largest freshwater fishery of the world.
The issues associated with Asian carp show how population and community ecology, fisheries management, and politics intersect on issues of vital importance to the human food supply and economy. Socio-political issues like this make extensive use of the sciences of population ecology (the study of members of a particular species occupying a particular area known as a habitat) and community ecology (the study of the interaction of all species within a habitat).
Community Dynamics
Community dynamics are the changes in community structure and composition over time. Sometimes these changes are induced by environmental disturbances such as volcanoes, earthquakes, storms, fires, and climate change. Communities with a stable structure are said to be at equilibrium. Following a disturbance, the community may or may not return to the equilibrium state.
Succession describes the sequential appearance and disappearance of species in a community over time. In primary succession, newly exposed or newly formed land is colonized by living things; in secondary succession, part of an ecosystem is disturbed and remnants of the previous community remain.
Primary Succession and Pioneer Species
Primary succession occurs when new land is formed or rock is exposed: for example, following the eruption of volcanoes, such as those on the Big Island of Hawaii. As lava flows into the ocean, new land is continually being formed. On the Big Island, approximately 32 acres of land is added each year. First, weathering and other natural forces break down the substrate enough for the establishment of certain hearty plants and lichens with few soil requirements, known as pioneer species (Figure \(15\)). These species help to further break down the mineral rich lava into soil where other, less hardy species will grow and eventually replace the pioneer species. In addition, as these early species grow and die, they add to an ever-growing layer of decomposing organic material and contribute to soil formation. Over time the area will reach an equilibrium state, with a set of organisms quite different from the pioneer species.
Secondary succession
A classic example of secondary succession occurs in oak and hickory forests cleared by wildfire (Figure \(16\)). Wildfires will burn most vegetation and kill those animals unable to flee the area. Their nutrients, however, are returned to the ground in the form of ash. Thus, even when areas are devoid of life due to severe fires, the area will soon be ready for new life to take hold.
Before the fire, the vegetation was dominated by tall trees with access to the major plant energy resource: sunlight. Their height gave them access to sunlight while also shading the ground and other low-lying species. After the fire, though, these trees are no longer dominant. Thus, the first plants to grow back are usually annual plants followed within a few years by quickly growing and spreading grasses and other pioneer species. Due to, at least in part, changes in the environment brought on by the growth of the grasses and other species, over many years, shrubs will emerge along with small pine, oak, and hickory trees. These organisms are called intermediate species. Eventually, over 150 years, the forest will reach its equilibrium point where species composition is no longer changing and resembles the community before the fire. This equilibrium state is referred to as the climax community, which will remain stable until the next disturbance.
Summary
Communities include all the different species living in a given area. The variety of these species is called species richness. Many organisms have developed defenses against predation and herbivory, including mechanical defenses, warning coloration, and mimicry, as a result of evolution and the interaction with other members of the community. Two species cannot exist in the same habitat competing directly for the same resources. Species may form symbiotic relationships such as commensalism or mutualism. Community structure is described by its foundation and keystone species. Communities respond to environmental disturbances by succession (the predictable appearance of different types of plant species) until a stable community structure is established.
Glossary
aposematic coloration
warning coloration used as a defensive mechanism against predation
Batesian mimicry
type of mimicry where a non-harmful species takes on the warning colorations of a harmful one
camouflage
avoid detection by blending in with the background.
climax community
final stage of succession, where a stable community is formed by a characteristic assortment of plant and animal species
commensalism
relationship between species wherein one species benefits from the close, prolonged interaction, while the other species neither benefits nor is harmed
competitive exclusion principle
no two species within a habitat can coexist when they compete for the same resources at the same place and time
Emsleyan/Mertensian mimicry
type of mimicry where a harmful species resembles a less harmful one
environmental disturbance
change in the environment caused by natural disasters or human activities
foundation species
species which often forms the major structural portion of the habitat
host
organism a parasite lives on
island biogeography
study of life on island chains and how their geography interacts with the diversity of species found there
keystone species
species whose presence is key to maintaining biodiversity in an ecosystem and to upholding an ecological community’s structure
Müllerian mimicry
type of mimicry where species share warning coloration and all are harmful to predators
mutualism
symbiotic relationship between two species where both species benefit
parasite
organism that uses resources from another species, the host
pioneer species
first species to appear in primary and secondary succession
primary succession
succession on land that previously has had no life
relative species abundance
absolute population size of a particular species relative to the population sizes of other species within the community
secondary succession
succession in response to environmental disturbances that move a community away from its equilibrium
species richness
number of different species in a community
symbiosis
close interaction between individuals of different species over an extended period of time that impacts the abundance and distribution of the associating populations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/55%3A_Community_Ecology/55.02%3A_The_Ecological_Niche_Concept/55.2.01%3A_Community_Ecology.txt |
Skills to Develop
• Discuss the predator-prey cycle
• Give examples of defenses against predation and herbivory
• Describe the competitive exclusion principle
• Give examples of symbiotic relationships between species
• Describe community structure and succession
Populations rarely, if ever, live in isolation from populations of other species. In most cases, numerous species share a habitat. The interactions between these populations play a major role in regulating population growth and abundance. All populations occupying the same habitat form a community: populations inhabiting a specific area at the same time. The number of species occupying the same habitat and their relative abundance is known as species diversity. Areas with low diversity, such as the glaciers of Antarctica, still contain a wide variety of living things, whereas the diversity of tropical rainforests is so great that it cannot be counted. Ecology is studied at the community level to understand how species interact with each other and compete for the same resources.
Predation and Herbivory
Perhaps the classical example of species interaction is predation: the hunting of prey by its predator. Nature shows on television highlight the drama of one living organism killing another. Populations of predators and prey in a community are not constant over time: in most cases, they vary in cycles that appear to be related. The most often cited example of predator-prey dynamics is seen in the cycling of the lynx (predator) and the snowshoe hare (prey), using nearly 200 year-old trapping data from North American forests (Figure \(1\)). This cycle of predator and prey lasts approximately 10 years, with the predator population lagging 1–2 years behind that of the prey population. As the hare numbers increase, there is more food available for the lynx, allowing the lynx population to increase as well. When the lynx population grows to a threshold level, however, they kill so many hares that hare population begins to decline, followed by a decline in the lynx population because of scarcity of food. When the lynx population is low, the hare population size begins to increase due, at least in part, to low predation pressure, starting the cycle anew.
The idea that the population cycling of the two species is entirely controlled by predation models has come under question. More recent studies have pointed to undefined density-dependent factors as being important in the cycling, in addition to predation. One possibility is that the cycling is inherent in the hare population due to density-dependent effects such as lower fecundity (maternal stress) caused by crowding when the hare population gets too dense. The hare cycling would then induce the cycling of the lynx because it is the lynxes’ major food source. The more we study communities, the more complexities we find, allowing ecologists to derive more accurate and sophisticated models of population dynamics.
Herbivory describes the consumption of plants by insects and other animals, and it is another interspecific relationship that affects populations. Unlike animals, most plants cannot outrun predators or use mimicry to hide from hungry animals. Some plants have developed mechanisms to defend against herbivory. Other species have developed mutualistic relationships; for example, herbivory provides a mechanism of seed distribution that aids in plant reproduction.
Defense Mechanisms against Predation and Herbivory
The study of communities must consider evolutionary forces that act on the members of the various populations contained within it. Species are not static, but slowly changing and adapting to their environment by natural selection and other evolutionary forces. Species have evolved numerous mechanisms to escape predation and herbivory. These defenses may be mechanical, chemical, physical, or behavioral.
Mechanical defenses, such as the presence of thorns on plants or the hard shell on turtles, discourage animal predation and herbivory by causing physical pain to the predator or by physically preventing the predator from being able to eat the prey. Chemical defenses are produced by many animals as well as plants, such as the foxglove which is extremely toxic when eaten. Figure \(2\) shows some organisms’ defenses against predation and herbivory.
Many species use their body shape and coloration to avoid being detected by predators. The tropical walking stick is an insect with the coloration and body shape of a twig which makes it very hard to see when stationary against a background of real twigs (Figure \(3\)a). In another example, the chameleon can change its color to match its surroundings (Figure \(3\)b). Both of these are examples of camouflage, or avoiding detection by blending in with the background.
Some species use coloration as a way of warning predators that they are not good to eat. For example, the cinnabar moth caterpillar, the fire-bellied toad, and many species of beetle have bright colors that warn of a foul taste, the presence of toxic chemical, and/or the ability to sting or bite, respectively. Predators that ignore this coloration and eat the organisms will experience their unpleasant taste or presence of toxic chemicals and learn not to eat them in the future. This type of defensive mechanism is called aposematic coloration, or warning coloration (Figure \(4\)).
While some predators learn to avoid eating certain potential prey because of their coloration, other species have evolved mechanisms to mimic this coloration to avoid being eaten, even though they themselves may not be unpleasant to eat or contain toxic chemicals. In Batesian mimicry, a harmless species imitates the warning coloration of a harmful one. Assuming they share the same predators, this coloration then protects the harmless ones, even though they do not have the same level of physical or chemical defenses against predation as the organism they mimic. Many insect species mimic the coloration of wasps or bees, which are stinging, venomous insects, thereby discouraging predation (Figure \(5\)).
In Müllerian mimicry, multiple species share the same warning coloration, but all of them actually have defenses. Figure \(6\) shows a variety of foul-tasting butterflies with similar coloration. In Emsleyan/Mertensian mimicry, a deadly prey mimics a less dangerous one, such as the venomous coral snake mimicking the non-venomous milk snake. This type of mimicry is extremely rare and more difficult to understand than the previous two types. For this type of mimicry to work, it is essential that eating the milk snake has unpleasant but not fatal consequences. Then, these predators learn not to eat snakes with this coloration, protecting the coral snake as well. If the snake were fatal to the predator, there would be no opportunity for the predator to learn not to eat it, and the benefit for the less toxic species would disappear.
Link to Learning
Go to this website to view stunning examples of mimicry.
Competitive Exclusion Principle
Resources are often limited within a habitat and multiple species may compete to obtain them. All species have an ecological niche in the ecosystem, which describes how they acquire the resources they need and how they interact with other species in the community. The competitive exclusion principle states that two species cannot occupy the same niche in a habitat. In other words, different species cannot coexist in a community if they are competing for all the same resources. An example of this principle is shown in Figure \(7\), with two protozoan species, Paramecium aurelia and Paramecium caudatum. When grown individually in the laboratory, they both thrive. But when they are placed together in the same test tube (habitat), P. aurelia outcompetes P. caudatum for food, leading to the latter’s eventual extinction.
This exclusion may be avoided if a population evolves to make use of a different resource, a different area of the habitat, or feeds during a different time of day, called resource partitioning. The two organisms are then said to occupy different microniches. These organisms coexist by minimizing direct competition.
Symbiosis
Symbiotic relationships, or symbioses (plural), are close interactions between individuals of different species over an extended period of time which impact the abundance and distribution of the associating populations. Most scientists accept this definition, but some restrict the term to only those species that are mutualistic, where both individuals benefit from the interaction. In this discussion, the broader definition will be used.
Commensalism
A commensal relationship occurs when one species benefits from the close, prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure \(8\)). The tree is not harmed by the presence of the nest among its branches. The nests are light and produce little strain on the structural integrity of the branch, and most of the leaves, which the tree uses to get energy by photosynthesis, are above the nest so they are unaffected. The bird, on the other hand, benefits greatly. If the bird had to nest in the open, its eggs and young would be vulnerable to predators. Another example of a commensal relationship is the clown fish and the sea anemone. The sea anemone is not harmed by the fish, and the fish benefits with protection from predators who would be stung upon nearing the sea anemone.
Mutualism
A second type of symbiotic relationship is called mutualism, where two species benefit from their interaction. Some scientists believe that these are the only true examples of symbiosis. For example, termites have a mutualistic relationship with protozoa that live in the insect’s gut (Figure \(9\)a). The termite benefits from the ability of bacterial symbionts within the protozoa to digest cellulose. The termite itself cannot do this, and without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa and the bacterial symbionts benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. Lichens have a mutualistic relationship between fungus and photosynthetic algae or bacteria (Figure \(9\)b). As these symbionts grow together, the glucose produced by the algae provides nourishment for both organisms, whereas the physical structure of the lichen protects the algae from the elements and makes certain nutrients in the atmosphere more available to the algae.
Parasitism
A parasite is an organism that lives in or on another living organism and derives nutrients from it. In this relationship, the parasite benefits, but the organism being fed upon, the host, is harmed. The host is usually weakened by the parasite as it siphons resources the host would normally use to maintain itself. The parasite, however, is unlikely to kill the host, especially not quickly, because this would allow no time for the organism to complete its reproductive cycle by spreading to another host.
The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm is a parasite that causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure \(10\)). The tapeworm can live inside the intestine of the host for several years, benefiting from the food the host is bringing into its gut by eating, and may grow to be over 50 ft long by adding segments. The parasite moves from species to species in a cycle, making two hosts necessary to complete its life cycle. Another common parasite is Plasmodium falciparum, the protozoan cause of malaria, a significant disease in many parts of the world. Living in human liver and red blood cells, the organism reproduces asexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to human by mosquitoes, one of many arthropod-borne infectious diseases.
Characteristics of Communities
Communities are complex entities that can be characterized by their structure (the types and numbers of species present) and dynamics (how communities change over time). Understanding community structure and dynamics enables community ecologists to manage ecosystems more effectively.
Foundation Species
Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are usually the primary producers: organisms that bring most of the energy into the community. Kelp, brown algae, is a foundation species, forming the basis of the kelp forests off the coast of California.
Foundation species may physically modify the environment to produce and maintain habitats that benefit the other organisms that use them. An example is the photosynthetic corals of the coral reef (Figure \(11\)). Corals themselves are not photosynthetic, but harbor symbionts within their body tissues (dinoflagellates called zooxanthellae) that perform photosynthesis; this is another example of a mutualism. The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.
Biodiversity, Species Richness, and Relative Species Abundance
Biodiversity describes a community’s biological complexity: it is measured by the number of different species (species richness) in a particular area and their relative abundance (species evenness). The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term that is used to describe the number of species living in a habitat or biome. Species richness varies across the globe (Figure \(12\)). One factor in determining species richness is latitude, with the greatest species richness occurring in ecosystems near the equator, which often have warmer temperatures, large amounts of rainfall, and low seasonality. The lowest species richness occurs near the poles, which are much colder, drier, and thus less conducive to life in Geologic time (time since glaciations). The predictability of climate or productivity is also an important factor. Other factors influence species richness as well. For example, the study of island biogeography attempts to explain the relatively high species richness found in certain isolated island chains, including the Galápagos Islands that inspired the young Darwin. Relative species abundance is the number of individuals in a species relative to the total number of individuals in all species within a habitat, ecosystem, or biome. Foundation species often have the highest relative abundance of species.
Keystone Species
A keystone species is one whose presence is key to maintaining biodiversity within an ecosystem and to upholding an ecological community’s structure. The intertidal sea star, Pisaster ochraceus, of the northwestern United States is a keystone species (Figure \(13\)). Studies have shown that when this organism is removed from communities, populations of their natural prey (mussels) increase, completely altering the species composition and reducing biodiversity. Another keystone species is the banded tetra, a fish in tropical streams, which supplies nearly all of the phosphorus, a necessary inorganic nutrient, to the rest of the community. If these fish were to become extinct, the community would be greatly affected.
Everyday Connection: Invasive Species
Invasive species are non-native organisms that, when introduced to an area out of their native range, threaten the ecosystem balance of that habitat. Many such species exist in the United States, as shown in Figure \(14\). Whether enjoying a forest hike, taking a summer boat trip, or simply walking down an urban street, you have likely encountered an invasive species.
One of the many recent proliferations of an invasive species concerns the growth of Asian carp populations. Asian carp were introduced to the United States in the 1970s by fisheries and sewage treatment facilities that used the fish’s excellent filter feeding capabilities to clean their ponds of excess plankton. Some of the fish escaped, however, and by the 1980s they had colonized many waterways of the Mississippi River basin, including the Illinois and Missouri Rivers.
Voracious eaters and rapid reproducers, Asian carp may outcompete native species for food, potentially leading to their extinction. For example, black carp are voracious eaters of native mussels and snails, limiting this food source for native fish species. Silver carp eat plankton that native mussels and snails feed on, reducing this food source by a different alteration of the food web. In some areas of the Mississippi River, Asian carp species have become the most predominant, effectively outcompeting native fishes for habitat. In some parts of the Illinois River, Asian carp constitute 95 percent of the community's biomass. Although edible, the fish is bony and not a desired food in the United States. Moreover, their presence threatens the native fish and fisheries of the Great Lakes, which are important to local economies and recreational anglers. Asian carp have even injured humans. The fish, frightened by the sound of approaching motorboats, thrust themselves into the air, often landing in the boat or directly hitting the boaters.
The Great Lakes and their prized salmon and lake trout fisheries are also being threatened by these invasive fish. Asian carp have already colonized rivers and canals that lead into Lake Michigan. One infested waterway of particular importance is the Chicago Sanitary and Ship Channel, the major supply waterway linking the Great Lakes to the Mississippi River. To prevent the Asian carp from leaving the canal, a series of electric barriers have been successfully used to discourage their migration; however, the threat is significant enough that several states and Canada have sued to have the Chicago channel permanently cut off from Lake Michigan. Local and national politicians have weighed in on how to solve the problem, but no one knows whether the Asian carp will ultimately be considered a nuisance, like other invasive species such as the water hyacinth and zebra mussel, or whether it will be the destroyer of the largest freshwater fishery of the world.
The issues associated with Asian carp show how population and community ecology, fisheries management, and politics intersect on issues of vital importance to the human food supply and economy. Socio-political issues like this make extensive use of the sciences of population ecology (the study of members of a particular species occupying a particular area known as a habitat) and community ecology (the study of the interaction of all species within a habitat).
Community Dynamics
Community dynamics are the changes in community structure and composition over time. Sometimes these changes are induced by environmental disturbances such as volcanoes, earthquakes, storms, fires, and climate change. Communities with a stable structure are said to be at equilibrium. Following a disturbance, the community may or may not return to the equilibrium state.
Succession describes the sequential appearance and disappearance of species in a community over time. In primary succession, newly exposed or newly formed land is colonized by living things; in secondary succession, part of an ecosystem is disturbed and remnants of the previous community remain.
Primary Succession and Pioneer Species
Primary succession occurs when new land is formed or rock is exposed: for example, following the eruption of volcanoes, such as those on the Big Island of Hawaii. As lava flows into the ocean, new land is continually being formed. On the Big Island, approximately 32 acres of land is added each year. First, weathering and other natural forces break down the substrate enough for the establishment of certain hearty plants and lichens with few soil requirements, known as pioneer species (Figure \(15\)). These species help to further break down the mineral rich lava into soil where other, less hardy species will grow and eventually replace the pioneer species. In addition, as these early species grow and die, they add to an ever-growing layer of decomposing organic material and contribute to soil formation. Over time the area will reach an equilibrium state, with a set of organisms quite different from the pioneer species.
Secondary succession
A classic example of secondary succession occurs in oak and hickory forests cleared by wildfire (Figure \(16\)). Wildfires will burn most vegetation and kill those animals unable to flee the area. Their nutrients, however, are returned to the ground in the form of ash. Thus, even when areas are devoid of life due to severe fires, the area will soon be ready for new life to take hold.
Before the fire, the vegetation was dominated by tall trees with access to the major plant energy resource: sunlight. Their height gave them access to sunlight while also shading the ground and other low-lying species. After the fire, though, these trees are no longer dominant. Thus, the first plants to grow back are usually annual plants followed within a few years by quickly growing and spreading grasses and other pioneer species. Due to, at least in part, changes in the environment brought on by the growth of the grasses and other species, over many years, shrubs will emerge along with small pine, oak, and hickory trees. These organisms are called intermediate species. Eventually, over 150 years, the forest will reach its equilibrium point where species composition is no longer changing and resembles the community before the fire. This equilibrium state is referred to as the climax community, which will remain stable until the next disturbance.
Summary
Communities include all the different species living in a given area. The variety of these species is called species richness. Many organisms have developed defenses against predation and herbivory, including mechanical defenses, warning coloration, and mimicry, as a result of evolution and the interaction with other members of the community. Two species cannot exist in the same habitat competing directly for the same resources. Species may form symbiotic relationships such as commensalism or mutualism. Community structure is described by its foundation and keystone species. Communities respond to environmental disturbances by succession (the predictable appearance of different types of plant species) until a stable community structure is established.
Glossary
aposematic coloration
warning coloration used as a defensive mechanism against predation
Batesian mimicry
type of mimicry where a non-harmful species takes on the warning colorations of a harmful one
camouflage
avoid detection by blending in with the background.
climax community
final stage of succession, where a stable community is formed by a characteristic assortment of plant and animal species
commensalism
relationship between species wherein one species benefits from the close, prolonged interaction, while the other species neither benefits nor is harmed
competitive exclusion principle
no two species within a habitat can coexist when they compete for the same resources at the same place and time
Emsleyan/Mertensian mimicry
type of mimicry where a harmful species resembles a less harmful one
environmental disturbance
change in the environment caused by natural disasters or human activities
foundation species
species which often forms the major structural portion of the habitat
host
organism a parasite lives on
island biogeography
study of life on island chains and how their geography interacts with the diversity of species found there
keystone species
species whose presence is key to maintaining biodiversity in an ecosystem and to upholding an ecological community’s structure
Müllerian mimicry
type of mimicry where species share warning coloration and all are harmful to predators
mutualism
symbiotic relationship between two species where both species benefit
parasite
organism that uses resources from another species, the host
pioneer species
first species to appear in primary and secondary succession
primary succession
succession on land that previously has had no life
relative species abundance
absolute population size of a particular species relative to the population sizes of other species within the community
secondary succession
succession in response to environmental disturbances that move a community away from its equilibrium
species richness
number of different species in a community
symbiosis
close interaction between individuals of different species over an extended period of time that impacts the abundance and distribution of the associating populations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/55%3A_Community_Ecology/55.03%3A_Predator-Prey_Relationships/55.3.01%3A_Community_Ecology.txt |
Skills to Develop
• Discuss the predator-prey cycle
• Give examples of defenses against predation and herbivory
• Describe the competitive exclusion principle
• Give examples of symbiotic relationships between species
• Describe community structure and succession
Populations rarely, if ever, live in isolation from populations of other species. In most cases, numerous species share a habitat. The interactions between these populations play a major role in regulating population growth and abundance. All populations occupying the same habitat form a community: populations inhabiting a specific area at the same time. The number of species occupying the same habitat and their relative abundance is known as species diversity. Areas with low diversity, such as the glaciers of Antarctica, still contain a wide variety of living things, whereas the diversity of tropical rainforests is so great that it cannot be counted. Ecology is studied at the community level to understand how species interact with each other and compete for the same resources.
Predation and Herbivory
Perhaps the classical example of species interaction is predation: the hunting of prey by its predator. Nature shows on television highlight the drama of one living organism killing another. Populations of predators and prey in a community are not constant over time: in most cases, they vary in cycles that appear to be related. The most often cited example of predator-prey dynamics is seen in the cycling of the lynx (predator) and the snowshoe hare (prey), using nearly 200 year-old trapping data from North American forests (Figure \(1\)). This cycle of predator and prey lasts approximately 10 years, with the predator population lagging 1–2 years behind that of the prey population. As the hare numbers increase, there is more food available for the lynx, allowing the lynx population to increase as well. When the lynx population grows to a threshold level, however, they kill so many hares that hare population begins to decline, followed by a decline in the lynx population because of scarcity of food. When the lynx population is low, the hare population size begins to increase due, at least in part, to low predation pressure, starting the cycle anew.
The idea that the population cycling of the two species is entirely controlled by predation models has come under question. More recent studies have pointed to undefined density-dependent factors as being important in the cycling, in addition to predation. One possibility is that the cycling is inherent in the hare population due to density-dependent effects such as lower fecundity (maternal stress) caused by crowding when the hare population gets too dense. The hare cycling would then induce the cycling of the lynx because it is the lynxes’ major food source. The more we study communities, the more complexities we find, allowing ecologists to derive more accurate and sophisticated models of population dynamics.
Herbivory describes the consumption of plants by insects and other animals, and it is another interspecific relationship that affects populations. Unlike animals, most plants cannot outrun predators or use mimicry to hide from hungry animals. Some plants have developed mechanisms to defend against herbivory. Other species have developed mutualistic relationships; for example, herbivory provides a mechanism of seed distribution that aids in plant reproduction.
Defense Mechanisms against Predation and Herbivory
The study of communities must consider evolutionary forces that act on the members of the various populations contained within it. Species are not static, but slowly changing and adapting to their environment by natural selection and other evolutionary forces. Species have evolved numerous mechanisms to escape predation and herbivory. These defenses may be mechanical, chemical, physical, or behavioral.
Mechanical defenses, such as the presence of thorns on plants or the hard shell on turtles, discourage animal predation and herbivory by causing physical pain to the predator or by physically preventing the predator from being able to eat the prey. Chemical defenses are produced by many animals as well as plants, such as the foxglove which is extremely toxic when eaten. Figure \(2\) shows some organisms’ defenses against predation and herbivory.
Many species use their body shape and coloration to avoid being detected by predators. The tropical walking stick is an insect with the coloration and body shape of a twig which makes it very hard to see when stationary against a background of real twigs (Figure \(3\)a). In another example, the chameleon can change its color to match its surroundings (Figure \(3\)b). Both of these are examples of camouflage, or avoiding detection by blending in with the background.
Some species use coloration as a way of warning predators that they are not good to eat. For example, the cinnabar moth caterpillar, the fire-bellied toad, and many species of beetle have bright colors that warn of a foul taste, the presence of toxic chemical, and/or the ability to sting or bite, respectively. Predators that ignore this coloration and eat the organisms will experience their unpleasant taste or presence of toxic chemicals and learn not to eat them in the future. This type of defensive mechanism is called aposematic coloration, or warning coloration (Figure \(4\)).
While some predators learn to avoid eating certain potential prey because of their coloration, other species have evolved mechanisms to mimic this coloration to avoid being eaten, even though they themselves may not be unpleasant to eat or contain toxic chemicals. In Batesian mimicry, a harmless species imitates the warning coloration of a harmful one. Assuming they share the same predators, this coloration then protects the harmless ones, even though they do not have the same level of physical or chemical defenses against predation as the organism they mimic. Many insect species mimic the coloration of wasps or bees, which are stinging, venomous insects, thereby discouraging predation (Figure \(5\)).
In Müllerian mimicry, multiple species share the same warning coloration, but all of them actually have defenses. Figure \(6\) shows a variety of foul-tasting butterflies with similar coloration. In Emsleyan/Mertensian mimicry, a deadly prey mimics a less dangerous one, such as the venomous coral snake mimicking the non-venomous milk snake. This type of mimicry is extremely rare and more difficult to understand than the previous two types. For this type of mimicry to work, it is essential that eating the milk snake has unpleasant but not fatal consequences. Then, these predators learn not to eat snakes with this coloration, protecting the coral snake as well. If the snake were fatal to the predator, there would be no opportunity for the predator to learn not to eat it, and the benefit for the less toxic species would disappear.
Link to Learning
Go to this website to view stunning examples of mimicry.
Competitive Exclusion Principle
Resources are often limited within a habitat and multiple species may compete to obtain them. All species have an ecological niche in the ecosystem, which describes how they acquire the resources they need and how they interact with other species in the community. The competitive exclusion principle states that two species cannot occupy the same niche in a habitat. In other words, different species cannot coexist in a community if they are competing for all the same resources. An example of this principle is shown in Figure \(7\), with two protozoan species, Paramecium aurelia and Paramecium caudatum. When grown individually in the laboratory, they both thrive. But when they are placed together in the same test tube (habitat), P. aurelia outcompetes P. caudatum for food, leading to the latter’s eventual extinction.
This exclusion may be avoided if a population evolves to make use of a different resource, a different area of the habitat, or feeds during a different time of day, called resource partitioning. The two organisms are then said to occupy different microniches. These organisms coexist by minimizing direct competition.
Symbiosis
Symbiotic relationships, or symbioses (plural), are close interactions between individuals of different species over an extended period of time which impact the abundance and distribution of the associating populations. Most scientists accept this definition, but some restrict the term to only those species that are mutualistic, where both individuals benefit from the interaction. In this discussion, the broader definition will be used.
Commensalism
A commensal relationship occurs when one species benefits from the close, prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure \(8\)). The tree is not harmed by the presence of the nest among its branches. The nests are light and produce little strain on the structural integrity of the branch, and most of the leaves, which the tree uses to get energy by photosynthesis, are above the nest so they are unaffected. The bird, on the other hand, benefits greatly. If the bird had to nest in the open, its eggs and young would be vulnerable to predators. Another example of a commensal relationship is the clown fish and the sea anemone. The sea anemone is not harmed by the fish, and the fish benefits with protection from predators who would be stung upon nearing the sea anemone.
Mutualism
A second type of symbiotic relationship is called mutualism, where two species benefit from their interaction. Some scientists believe that these are the only true examples of symbiosis. For example, termites have a mutualistic relationship with protozoa that live in the insect’s gut (Figure \(9\)a). The termite benefits from the ability of bacterial symbionts within the protozoa to digest cellulose. The termite itself cannot do this, and without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa and the bacterial symbionts benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. Lichens have a mutualistic relationship between fungus and photosynthetic algae or bacteria (Figure \(9\)b). As these symbionts grow together, the glucose produced by the algae provides nourishment for both organisms, whereas the physical structure of the lichen protects the algae from the elements and makes certain nutrients in the atmosphere more available to the algae.
Parasitism
A parasite is an organism that lives in or on another living organism and derives nutrients from it. In this relationship, the parasite benefits, but the organism being fed upon, the host, is harmed. The host is usually weakened by the parasite as it siphons resources the host would normally use to maintain itself. The parasite, however, is unlikely to kill the host, especially not quickly, because this would allow no time for the organism to complete its reproductive cycle by spreading to another host.
The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm is a parasite that causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure \(10\)). The tapeworm can live inside the intestine of the host for several years, benefiting from the food the host is bringing into its gut by eating, and may grow to be over 50 ft long by adding segments. The parasite moves from species to species in a cycle, making two hosts necessary to complete its life cycle. Another common parasite is Plasmodium falciparum, the protozoan cause of malaria, a significant disease in many parts of the world. Living in human liver and red blood cells, the organism reproduces asexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to human by mosquitoes, one of many arthropod-borne infectious diseases.
Characteristics of Communities
Communities are complex entities that can be characterized by their structure (the types and numbers of species present) and dynamics (how communities change over time). Understanding community structure and dynamics enables community ecologists to manage ecosystems more effectively.
Foundation Species
Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are usually the primary producers: organisms that bring most of the energy into the community. Kelp, brown algae, is a foundation species, forming the basis of the kelp forests off the coast of California.
Foundation species may physically modify the environment to produce and maintain habitats that benefit the other organisms that use them. An example is the photosynthetic corals of the coral reef (Figure \(11\)). Corals themselves are not photosynthetic, but harbor symbionts within their body tissues (dinoflagellates called zooxanthellae) that perform photosynthesis; this is another example of a mutualism. The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.
Biodiversity, Species Richness, and Relative Species Abundance
Biodiversity describes a community’s biological complexity: it is measured by the number of different species (species richness) in a particular area and their relative abundance (species evenness). The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term that is used to describe the number of species living in a habitat or biome. Species richness varies across the globe (Figure \(12\)). One factor in determining species richness is latitude, with the greatest species richness occurring in ecosystems near the equator, which often have warmer temperatures, large amounts of rainfall, and low seasonality. The lowest species richness occurs near the poles, which are much colder, drier, and thus less conducive to life in Geologic time (time since glaciations). The predictability of climate or productivity is also an important factor. Other factors influence species richness as well. For example, the study of island biogeography attempts to explain the relatively high species richness found in certain isolated island chains, including the Galápagos Islands that inspired the young Darwin. Relative species abundance is the number of individuals in a species relative to the total number of individuals in all species within a habitat, ecosystem, or biome. Foundation species often have the highest relative abundance of species.
Keystone Species
A keystone species is one whose presence is key to maintaining biodiversity within an ecosystem and to upholding an ecological community’s structure. The intertidal sea star, Pisaster ochraceus, of the northwestern United States is a keystone species (Figure \(13\)). Studies have shown that when this organism is removed from communities, populations of their natural prey (mussels) increase, completely altering the species composition and reducing biodiversity. Another keystone species is the banded tetra, a fish in tropical streams, which supplies nearly all of the phosphorus, a necessary inorganic nutrient, to the rest of the community. If these fish were to become extinct, the community would be greatly affected.
Everyday Connection: Invasive Species
Invasive species are non-native organisms that, when introduced to an area out of their native range, threaten the ecosystem balance of that habitat. Many such species exist in the United States, as shown in Figure \(14\). Whether enjoying a forest hike, taking a summer boat trip, or simply walking down an urban street, you have likely encountered an invasive species.
One of the many recent proliferations of an invasive species concerns the growth of Asian carp populations. Asian carp were introduced to the United States in the 1970s by fisheries and sewage treatment facilities that used the fish’s excellent filter feeding capabilities to clean their ponds of excess plankton. Some of the fish escaped, however, and by the 1980s they had colonized many waterways of the Mississippi River basin, including the Illinois and Missouri Rivers.
Voracious eaters and rapid reproducers, Asian carp may outcompete native species for food, potentially leading to their extinction. For example, black carp are voracious eaters of native mussels and snails, limiting this food source for native fish species. Silver carp eat plankton that native mussels and snails feed on, reducing this food source by a different alteration of the food web. In some areas of the Mississippi River, Asian carp species have become the most predominant, effectively outcompeting native fishes for habitat. In some parts of the Illinois River, Asian carp constitute 95 percent of the community's biomass. Although edible, the fish is bony and not a desired food in the United States. Moreover, their presence threatens the native fish and fisheries of the Great Lakes, which are important to local economies and recreational anglers. Asian carp have even injured humans. The fish, frightened by the sound of approaching motorboats, thrust themselves into the air, often landing in the boat or directly hitting the boaters.
The Great Lakes and their prized salmon and lake trout fisheries are also being threatened by these invasive fish. Asian carp have already colonized rivers and canals that lead into Lake Michigan. One infested waterway of particular importance is the Chicago Sanitary and Ship Channel, the major supply waterway linking the Great Lakes to the Mississippi River. To prevent the Asian carp from leaving the canal, a series of electric barriers have been successfully used to discourage their migration; however, the threat is significant enough that several states and Canada have sued to have the Chicago channel permanently cut off from Lake Michigan. Local and national politicians have weighed in on how to solve the problem, but no one knows whether the Asian carp will ultimately be considered a nuisance, like other invasive species such as the water hyacinth and zebra mussel, or whether it will be the destroyer of the largest freshwater fishery of the world.
The issues associated with Asian carp show how population and community ecology, fisheries management, and politics intersect on issues of vital importance to the human food supply and economy. Socio-political issues like this make extensive use of the sciences of population ecology (the study of members of a particular species occupying a particular area known as a habitat) and community ecology (the study of the interaction of all species within a habitat).
Community Dynamics
Community dynamics are the changes in community structure and composition over time. Sometimes these changes are induced by environmental disturbances such as volcanoes, earthquakes, storms, fires, and climate change. Communities with a stable structure are said to be at equilibrium. Following a disturbance, the community may or may not return to the equilibrium state.
Succession describes the sequential appearance and disappearance of species in a community over time. In primary succession, newly exposed or newly formed land is colonized by living things; in secondary succession, part of an ecosystem is disturbed and remnants of the previous community remain.
Primary Succession and Pioneer Species
Primary succession occurs when new land is formed or rock is exposed: for example, following the eruption of volcanoes, such as those on the Big Island of Hawaii. As lava flows into the ocean, new land is continually being formed. On the Big Island, approximately 32 acres of land is added each year. First, weathering and other natural forces break down the substrate enough for the establishment of certain hearty plants and lichens with few soil requirements, known as pioneer species (Figure \(15\)). These species help to further break down the mineral rich lava into soil where other, less hardy species will grow and eventually replace the pioneer species. In addition, as these early species grow and die, they add to an ever-growing layer of decomposing organic material and contribute to soil formation. Over time the area will reach an equilibrium state, with a set of organisms quite different from the pioneer species.
Secondary succession
A classic example of secondary succession occurs in oak and hickory forests cleared by wildfire (Figure \(16\)). Wildfires will burn most vegetation and kill those animals unable to flee the area. Their nutrients, however, are returned to the ground in the form of ash. Thus, even when areas are devoid of life due to severe fires, the area will soon be ready for new life to take hold.
Before the fire, the vegetation was dominated by tall trees with access to the major plant energy resource: sunlight. Their height gave them access to sunlight while also shading the ground and other low-lying species. After the fire, though, these trees are no longer dominant. Thus, the first plants to grow back are usually annual plants followed within a few years by quickly growing and spreading grasses and other pioneer species. Due to, at least in part, changes in the environment brought on by the growth of the grasses and other species, over many years, shrubs will emerge along with small pine, oak, and hickory trees. These organisms are called intermediate species. Eventually, over 150 years, the forest will reach its equilibrium point where species composition is no longer changing and resembles the community before the fire. This equilibrium state is referred to as the climax community, which will remain stable until the next disturbance.
Summary
Communities include all the different species living in a given area. The variety of these species is called species richness. Many organisms have developed defenses against predation and herbivory, including mechanical defenses, warning coloration, and mimicry, as a result of evolution and the interaction with other members of the community. Two species cannot exist in the same habitat competing directly for the same resources. Species may form symbiotic relationships such as commensalism or mutualism. Community structure is described by its foundation and keystone species. Communities respond to environmental disturbances by succession (the predictable appearance of different types of plant species) until a stable community structure is established.
Glossary
aposematic coloration
warning coloration used as a defensive mechanism against predation
Batesian mimicry
type of mimicry where a non-harmful species takes on the warning colorations of a harmful one
camouflage
avoid detection by blending in with the background.
climax community
final stage of succession, where a stable community is formed by a characteristic assortment of plant and animal species
commensalism
relationship between species wherein one species benefits from the close, prolonged interaction, while the other species neither benefits nor is harmed
competitive exclusion principle
no two species within a habitat can coexist when they compete for the same resources at the same place and time
Emsleyan/Mertensian mimicry
type of mimicry where a harmful species resembles a less harmful one
environmental disturbance
change in the environment caused by natural disasters or human activities
foundation species
species which often forms the major structural portion of the habitat
host
organism a parasite lives on
island biogeography
study of life on island chains and how their geography interacts with the diversity of species found there
keystone species
species whose presence is key to maintaining biodiversity in an ecosystem and to upholding an ecological community’s structure
Müllerian mimicry
type of mimicry where species share warning coloration and all are harmful to predators
mutualism
symbiotic relationship between two species where both species benefit
parasite
organism that uses resources from another species, the host
pioneer species
first species to appear in primary and secondary succession
primary succession
succession on land that previously has had no life
relative species abundance
absolute population size of a particular species relative to the population sizes of other species within the community
secondary succession
succession in response to environmental disturbances that move a community away from its equilibrium
species richness
number of different species in a community
symbiosis
close interaction between individuals of different species over an extended period of time that impacts the abundance and distribution of the associating populations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/55%3A_Community_Ecology/55.04%3A_The_Many_Types_of_Species_Interactions/55.4.01%3A_Community_Ecology.txt |
Skills to Develop
• Discuss the predator-prey cycle
• Give examples of defenses against predation and herbivory
• Describe the competitive exclusion principle
• Give examples of symbiotic relationships between species
• Describe community structure and succession
Populations rarely, if ever, live in isolation from populations of other species. In most cases, numerous species share a habitat. The interactions between these populations play a major role in regulating population growth and abundance. All populations occupying the same habitat form a community: populations inhabiting a specific area at the same time. The number of species occupying the same habitat and their relative abundance is known as species diversity. Areas with low diversity, such as the glaciers of Antarctica, still contain a wide variety of living things, whereas the diversity of tropical rainforests is so great that it cannot be counted. Ecology is studied at the community level to understand how species interact with each other and compete for the same resources.
Predation and Herbivory
Perhaps the classical example of species interaction is predation: the hunting of prey by its predator. Nature shows on television highlight the drama of one living organism killing another. Populations of predators and prey in a community are not constant over time: in most cases, they vary in cycles that appear to be related. The most often cited example of predator-prey dynamics is seen in the cycling of the lynx (predator) and the snowshoe hare (prey), using nearly 200 year-old trapping data from North American forests (Figure \(1\)). This cycle of predator and prey lasts approximately 10 years, with the predator population lagging 1–2 years behind that of the prey population. As the hare numbers increase, there is more food available for the lynx, allowing the lynx population to increase as well. When the lynx population grows to a threshold level, however, they kill so many hares that hare population begins to decline, followed by a decline in the lynx population because of scarcity of food. When the lynx population is low, the hare population size begins to increase due, at least in part, to low predation pressure, starting the cycle anew.
The idea that the population cycling of the two species is entirely controlled by predation models has come under question. More recent studies have pointed to undefined density-dependent factors as being important in the cycling, in addition to predation. One possibility is that the cycling is inherent in the hare population due to density-dependent effects such as lower fecundity (maternal stress) caused by crowding when the hare population gets too dense. The hare cycling would then induce the cycling of the lynx because it is the lynxes’ major food source. The more we study communities, the more complexities we find, allowing ecologists to derive more accurate and sophisticated models of population dynamics.
Herbivory describes the consumption of plants by insects and other animals, and it is another interspecific relationship that affects populations. Unlike animals, most plants cannot outrun predators or use mimicry to hide from hungry animals. Some plants have developed mechanisms to defend against herbivory. Other species have developed mutualistic relationships; for example, herbivory provides a mechanism of seed distribution that aids in plant reproduction.
Defense Mechanisms against Predation and Herbivory
The study of communities must consider evolutionary forces that act on the members of the various populations contained within it. Species are not static, but slowly changing and adapting to their environment by natural selection and other evolutionary forces. Species have evolved numerous mechanisms to escape predation and herbivory. These defenses may be mechanical, chemical, physical, or behavioral.
Mechanical defenses, such as the presence of thorns on plants or the hard shell on turtles, discourage animal predation and herbivory by causing physical pain to the predator or by physically preventing the predator from being able to eat the prey. Chemical defenses are produced by many animals as well as plants, such as the foxglove which is extremely toxic when eaten. Figure \(2\) shows some organisms’ defenses against predation and herbivory.
Many species use their body shape and coloration to avoid being detected by predators. The tropical walking stick is an insect with the coloration and body shape of a twig which makes it very hard to see when stationary against a background of real twigs (Figure \(3\)a). In another example, the chameleon can change its color to match its surroundings (Figure \(3\)b). Both of these are examples of camouflage, or avoiding detection by blending in with the background.
Some species use coloration as a way of warning predators that they are not good to eat. For example, the cinnabar moth caterpillar, the fire-bellied toad, and many species of beetle have bright colors that warn of a foul taste, the presence of toxic chemical, and/or the ability to sting or bite, respectively. Predators that ignore this coloration and eat the organisms will experience their unpleasant taste or presence of toxic chemicals and learn not to eat them in the future. This type of defensive mechanism is called aposematic coloration, or warning coloration (Figure \(4\)).
While some predators learn to avoid eating certain potential prey because of their coloration, other species have evolved mechanisms to mimic this coloration to avoid being eaten, even though they themselves may not be unpleasant to eat or contain toxic chemicals. In Batesian mimicry, a harmless species imitates the warning coloration of a harmful one. Assuming they share the same predators, this coloration then protects the harmless ones, even though they do not have the same level of physical or chemical defenses against predation as the organism they mimic. Many insect species mimic the coloration of wasps or bees, which are stinging, venomous insects, thereby discouraging predation (Figure \(5\)).
In Müllerian mimicry, multiple species share the same warning coloration, but all of them actually have defenses. Figure \(6\) shows a variety of foul-tasting butterflies with similar coloration. In Emsleyan/Mertensian mimicry, a deadly prey mimics a less dangerous one, such as the venomous coral snake mimicking the non-venomous milk snake. This type of mimicry is extremely rare and more difficult to understand than the previous two types. For this type of mimicry to work, it is essential that eating the milk snake has unpleasant but not fatal consequences. Then, these predators learn not to eat snakes with this coloration, protecting the coral snake as well. If the snake were fatal to the predator, there would be no opportunity for the predator to learn not to eat it, and the benefit for the less toxic species would disappear.
Link to Learning
Go to this website to view stunning examples of mimicry.
Competitive Exclusion Principle
Resources are often limited within a habitat and multiple species may compete to obtain them. All species have an ecological niche in the ecosystem, which describes how they acquire the resources they need and how they interact with other species in the community. The competitive exclusion principle states that two species cannot occupy the same niche in a habitat. In other words, different species cannot coexist in a community if they are competing for all the same resources. An example of this principle is shown in Figure \(7\), with two protozoan species, Paramecium aurelia and Paramecium caudatum. When grown individually in the laboratory, they both thrive. But when they are placed together in the same test tube (habitat), P. aurelia outcompetes P. caudatum for food, leading to the latter’s eventual extinction.
This exclusion may be avoided if a population evolves to make use of a different resource, a different area of the habitat, or feeds during a different time of day, called resource partitioning. The two organisms are then said to occupy different microniches. These organisms coexist by minimizing direct competition.
Symbiosis
Symbiotic relationships, or symbioses (plural), are close interactions between individuals of different species over an extended period of time which impact the abundance and distribution of the associating populations. Most scientists accept this definition, but some restrict the term to only those species that are mutualistic, where both individuals benefit from the interaction. In this discussion, the broader definition will be used.
Commensalism
A commensal relationship occurs when one species benefits from the close, prolonged interaction, while the other neither benefits nor is harmed. Birds nesting in trees provide an example of a commensal relationship (Figure \(8\)). The tree is not harmed by the presence of the nest among its branches. The nests are light and produce little strain on the structural integrity of the branch, and most of the leaves, which the tree uses to get energy by photosynthesis, are above the nest so they are unaffected. The bird, on the other hand, benefits greatly. If the bird had to nest in the open, its eggs and young would be vulnerable to predators. Another example of a commensal relationship is the clown fish and the sea anemone. The sea anemone is not harmed by the fish, and the fish benefits with protection from predators who would be stung upon nearing the sea anemone.
Mutualism
A second type of symbiotic relationship is called mutualism, where two species benefit from their interaction. Some scientists believe that these are the only true examples of symbiosis. For example, termites have a mutualistic relationship with protozoa that live in the insect’s gut (Figure \(9\)a). The termite benefits from the ability of bacterial symbionts within the protozoa to digest cellulose. The termite itself cannot do this, and without the protozoa, it would not be able to obtain energy from its food (cellulose from the wood it chews and eats). The protozoa and the bacterial symbionts benefit by having a protective environment and a constant supply of food from the wood chewing actions of the termite. Lichens have a mutualistic relationship between fungus and photosynthetic algae or bacteria (Figure \(9\)b). As these symbionts grow together, the glucose produced by the algae provides nourishment for both organisms, whereas the physical structure of the lichen protects the algae from the elements and makes certain nutrients in the atmosphere more available to the algae.
Parasitism
A parasite is an organism that lives in or on another living organism and derives nutrients from it. In this relationship, the parasite benefits, but the organism being fed upon, the host, is harmed. The host is usually weakened by the parasite as it siphons resources the host would normally use to maintain itself. The parasite, however, is unlikely to kill the host, especially not quickly, because this would allow no time for the organism to complete its reproductive cycle by spreading to another host.
The reproductive cycles of parasites are often very complex, sometimes requiring more than one host species. A tapeworm is a parasite that causes disease in humans when contaminated, undercooked meat such as pork, fish, or beef is consumed (Figure \(10\)). The tapeworm can live inside the intestine of the host for several years, benefiting from the food the host is bringing into its gut by eating, and may grow to be over 50 ft long by adding segments. The parasite moves from species to species in a cycle, making two hosts necessary to complete its life cycle. Another common parasite is Plasmodium falciparum, the protozoan cause of malaria, a significant disease in many parts of the world. Living in human liver and red blood cells, the organism reproduces asexually in the gut of blood-feeding mosquitoes to complete its life cycle. Thus malaria is spread from human to human by mosquitoes, one of many arthropod-borne infectious diseases.
Characteristics of Communities
Communities are complex entities that can be characterized by their structure (the types and numbers of species present) and dynamics (how communities change over time). Understanding community structure and dynamics enables community ecologists to manage ecosystems more effectively.
Foundation Species
Foundation species are considered the “base” or “bedrock” of a community, having the greatest influence on its overall structure. They are usually the primary producers: organisms that bring most of the energy into the community. Kelp, brown algae, is a foundation species, forming the basis of the kelp forests off the coast of California.
Foundation species may physically modify the environment to produce and maintain habitats that benefit the other organisms that use them. An example is the photosynthetic corals of the coral reef (Figure \(11\)). Corals themselves are not photosynthetic, but harbor symbionts within their body tissues (dinoflagellates called zooxanthellae) that perform photosynthesis; this is another example of a mutualism. The exoskeletons of living and dead coral make up most of the reef structure, which protects many other species from waves and ocean currents.
Biodiversity, Species Richness, and Relative Species Abundance
Biodiversity describes a community’s biological complexity: it is measured by the number of different species (species richness) in a particular area and their relative abundance (species evenness). The area in question could be a habitat, a biome, or the entire biosphere. Species richness is the term that is used to describe the number of species living in a habitat or biome. Species richness varies across the globe (Figure \(12\)). One factor in determining species richness is latitude, with the greatest species richness occurring in ecosystems near the equator, which often have warmer temperatures, large amounts of rainfall, and low seasonality. The lowest species richness occurs near the poles, which are much colder, drier, and thus less conducive to life in Geologic time (time since glaciations). The predictability of climate or productivity is also an important factor. Other factors influence species richness as well. For example, the study of island biogeography attempts to explain the relatively high species richness found in certain isolated island chains, including the Galápagos Islands that inspired the young Darwin. Relative species abundance is the number of individuals in a species relative to the total number of individuals in all species within a habitat, ecosystem, or biome. Foundation species often have the highest relative abundance of species.
Keystone Species
A keystone species is one whose presence is key to maintaining biodiversity within an ecosystem and to upholding an ecological community’s structure. The intertidal sea star, Pisaster ochraceus, of the northwestern United States is a keystone species (Figure \(13\)). Studies have shown that when this organism is removed from communities, populations of their natural prey (mussels) increase, completely altering the species composition and reducing biodiversity. Another keystone species is the banded tetra, a fish in tropical streams, which supplies nearly all of the phosphorus, a necessary inorganic nutrient, to the rest of the community. If these fish were to become extinct, the community would be greatly affected.
Everyday Connection: Invasive Species
Invasive species are non-native organisms that, when introduced to an area out of their native range, threaten the ecosystem balance of that habitat. Many such species exist in the United States, as shown in Figure \(14\). Whether enjoying a forest hike, taking a summer boat trip, or simply walking down an urban street, you have likely encountered an invasive species.
One of the many recent proliferations of an invasive species concerns the growth of Asian carp populations. Asian carp were introduced to the United States in the 1970s by fisheries and sewage treatment facilities that used the fish’s excellent filter feeding capabilities to clean their ponds of excess plankton. Some of the fish escaped, however, and by the 1980s they had colonized many waterways of the Mississippi River basin, including the Illinois and Missouri Rivers.
Voracious eaters and rapid reproducers, Asian carp may outcompete native species for food, potentially leading to their extinction. For example, black carp are voracious eaters of native mussels and snails, limiting this food source for native fish species. Silver carp eat plankton that native mussels and snails feed on, reducing this food source by a different alteration of the food web. In some areas of the Mississippi River, Asian carp species have become the most predominant, effectively outcompeting native fishes for habitat. In some parts of the Illinois River, Asian carp constitute 95 percent of the community's biomass. Although edible, the fish is bony and not a desired food in the United States. Moreover, their presence threatens the native fish and fisheries of the Great Lakes, which are important to local economies and recreational anglers. Asian carp have even injured humans. The fish, frightened by the sound of approaching motorboats, thrust themselves into the air, often landing in the boat or directly hitting the boaters.
The Great Lakes and their prized salmon and lake trout fisheries are also being threatened by these invasive fish. Asian carp have already colonized rivers and canals that lead into Lake Michigan. One infested waterway of particular importance is the Chicago Sanitary and Ship Channel, the major supply waterway linking the Great Lakes to the Mississippi River. To prevent the Asian carp from leaving the canal, a series of electric barriers have been successfully used to discourage their migration; however, the threat is significant enough that several states and Canada have sued to have the Chicago channel permanently cut off from Lake Michigan. Local and national politicians have weighed in on how to solve the problem, but no one knows whether the Asian carp will ultimately be considered a nuisance, like other invasive species such as the water hyacinth and zebra mussel, or whether it will be the destroyer of the largest freshwater fishery of the world.
The issues associated with Asian carp show how population and community ecology, fisheries management, and politics intersect on issues of vital importance to the human food supply and economy. Socio-political issues like this make extensive use of the sciences of population ecology (the study of members of a particular species occupying a particular area known as a habitat) and community ecology (the study of the interaction of all species within a habitat).
Community Dynamics
Community dynamics are the changes in community structure and composition over time. Sometimes these changes are induced by environmental disturbances such as volcanoes, earthquakes, storms, fires, and climate change. Communities with a stable structure are said to be at equilibrium. Following a disturbance, the community may or may not return to the equilibrium state.
Succession describes the sequential appearance and disappearance of species in a community over time. In primary succession, newly exposed or newly formed land is colonized by living things; in secondary succession, part of an ecosystem is disturbed and remnants of the previous community remain.
Primary Succession and Pioneer Species
Primary succession occurs when new land is formed or rock is exposed: for example, following the eruption of volcanoes, such as those on the Big Island of Hawaii. As lava flows into the ocean, new land is continually being formed. On the Big Island, approximately 32 acres of land is added each year. First, weathering and other natural forces break down the substrate enough for the establishment of certain hearty plants and lichens with few soil requirements, known as pioneer species (Figure \(15\)). These species help to further break down the mineral rich lava into soil where other, less hardy species will grow and eventually replace the pioneer species. In addition, as these early species grow and die, they add to an ever-growing layer of decomposing organic material and contribute to soil formation. Over time the area will reach an equilibrium state, with a set of organisms quite different from the pioneer species.
Secondary succession
A classic example of secondary succession occurs in oak and hickory forests cleared by wildfire (Figure \(16\)). Wildfires will burn most vegetation and kill those animals unable to flee the area. Their nutrients, however, are returned to the ground in the form of ash. Thus, even when areas are devoid of life due to severe fires, the area will soon be ready for new life to take hold.
Before the fire, the vegetation was dominated by tall trees with access to the major plant energy resource: sunlight. Their height gave them access to sunlight while also shading the ground and other low-lying species. After the fire, though, these trees are no longer dominant. Thus, the first plants to grow back are usually annual plants followed within a few years by quickly growing and spreading grasses and other pioneer species. Due to, at least in part, changes in the environment brought on by the growth of the grasses and other species, over many years, shrubs will emerge along with small pine, oak, and hickory trees. These organisms are called intermediate species. Eventually, over 150 years, the forest will reach its equilibrium point where species composition is no longer changing and resembles the community before the fire. This equilibrium state is referred to as the climax community, which will remain stable until the next disturbance.
Summary
Communities include all the different species living in a given area. The variety of these species is called species richness. Many organisms have developed defenses against predation and herbivory, including mechanical defenses, warning coloration, and mimicry, as a result of evolution and the interaction with other members of the community. Two species cannot exist in the same habitat competing directly for the same resources. Species may form symbiotic relationships such as commensalism or mutualism. Community structure is described by its foundation and keystone species. Communities respond to environmental disturbances by succession (the predictable appearance of different types of plant species) until a stable community structure is established.
Glossary
aposematic coloration
warning coloration used as a defensive mechanism against predation
Batesian mimicry
type of mimicry where a non-harmful species takes on the warning colorations of a harmful one
camouflage
avoid detection by blending in with the background.
climax community
final stage of succession, where a stable community is formed by a characteristic assortment of plant and animal species
commensalism
relationship between species wherein one species benefits from the close, prolonged interaction, while the other species neither benefits nor is harmed
competitive exclusion principle
no two species within a habitat can coexist when they compete for the same resources at the same place and time
Emsleyan/Mertensian mimicry
type of mimicry where a harmful species resembles a less harmful one
environmental disturbance
change in the environment caused by natural disasters or human activities
foundation species
species which often forms the major structural portion of the habitat
host
organism a parasite lives on
island biogeography
study of life on island chains and how their geography interacts with the diversity of species found there
keystone species
species whose presence is key to maintaining biodiversity in an ecosystem and to upholding an ecological community’s structure
Müllerian mimicry
type of mimicry where species share warning coloration and all are harmful to predators
mutualism
symbiotic relationship between two species where both species benefit
parasite
organism that uses resources from another species, the host
pioneer species
first species to appear in primary and secondary succession
primary succession
succession on land that previously has had no life
relative species abundance
absolute population size of a particular species relative to the population sizes of other species within the community
secondary succession
succession in response to environmental disturbances that move a community away from its equilibrium
species richness
number of different species in a community
symbiosis
close interaction between individuals of different species over an extended period of time that impacts the abundance and distribution of the associating populations | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/55%3A_Community_Ecology/55.05%3A_Ecological_Successions_Disturbance_and_Species_Richness/55.5.01%3A_Community_Ecology.txt |
• 56.1: Biogeochemical Cycles
The matter that makes up living organisms is conserved and recycled. The six most common elements associated with organic molecules—carbon, nitrogen, hydrogen, oxygen, phosphorus, and sulfur—take a variety of chemical forms and may exist for long periods in the atmosphere, on land, in water, or beneath the Earth’s surface. Geologic processes, such as weathering, erosion, water drainage, and the subduction of the continental plates, all play a role in this recycling of materials.
• 56.2: The Flow of Energy in Ecosystems
All living things require energy in one form or another. Energy is required by most complex metabolic pathways (often in the form of adenosine triphosphate, ATP), especially those responsible for building large molecules from smaller compounds, and life itself is an energy-driven process. Living organisms would not be able to assemble macromolecules (proteins, lipids, nucleic acids, and complex carbohydrates) from their monomeric subunits without a constant energy input.
• 56.3: Trophic Level Interactions
• 56.4: Biodiversity and Ecosystem Stability
• 56.5: Island Biogeography
56: Dynamics of Ecosystems
Skills to Develop
• Discuss the biogeochemical cycles of water, carbon, nitrogen, phosphorus, and sulfur
• Explain how human activities have impacted these cycles and the potential consequences for Earth
Energy flows directionally through ecosystems, entering as sunlight (or inorganic molecules for chemoautotrophs) and leaving as heat during the many transfers between trophic levels. However, the matter that makes up living organisms is conserved and recycled. The six most common elements associated with organic molecules—carbon, nitrogen, hydrogen, oxygen, phosphorus, and sulfur—take a variety of chemical forms and may exist for long periods in the atmosphere, on land, in water, or beneath the Earth’s surface. Geologic processes, such as weathering, erosion, water drainage, and the subduction of the continental plates, all play a role in this recycling of materials. Because geology and chemistry have major roles in the study of this process, the recycling of inorganic matter between living organisms and their environment is called a biogeochemical cycle.
Water contains hydrogen and oxygen, which is essential to all living processes. The hydrosphere is the area of the Earth where water movement and storage occurs: as liquid water on the surface and beneath the surface or frozen (rivers, lakes, oceans, groundwater, polar ice caps, and glaciers), and as water vapor in the atmosphere. Carbon is found in all organic macromolecules and is an important constituent of fossil fuels. Nitrogen is a major component of our nucleic acids and proteins and is critical to human agriculture. Phosphorus, a major component of nucleic acid (along with nitrogen), is one of the main ingredients in artificial fertilizers used in agriculture and their associated environmental impacts on our surface water. Sulfur, critical to the 3–D folding of proteins (as in disulfide binding), is released into the atmosphere by the burning of fossil fuels, such as coal.
The cycling of these elements is interconnected. For example, the movement of water is critical for the leaching of nitrogen and phosphate into rivers, lakes, and oceans. Furthermore, the ocean itself is a major reservoir for carbon. Thus, mineral nutrients are cycled, either rapidly or slowly, through the entire biosphere, from one living organism to another, and between the biotic and abiotic world.
The Water (Hydrologic) Cycle
Water is the basis of all living processes. The human body is more than 1/2 water and human cells are more than 70 percent water. Thus, most land animals need a supply of fresh water to survive. However, when examining the stores of water on Earth, 97.5 percent of it is non-potable salt water (Figure $1$). Of the remaining water, 99 percent is locked underground as water or as ice. Thus, less than 1 percent of fresh water is easily accessible from lakes and rivers. Many living things, such as plants, animals, and fungi, are dependent on the small amount of fresh surface water supply, a lack of which can have massive effects on ecosystem dynamics. Humans, of course, have developed technologies to increase water availability, such as digging wells to harvest groundwater, storing rainwater, and using desalination to obtain drinkable water from the ocean. Although this pursuit of drinkable water has been ongoing throughout human history, the supply of fresh water is still a major issue in modern times.
Water cycling is extremely important to ecosystem dynamics. Water has a major influence on climate and, thus, on the environments of ecosystems, some located on distant parts of the Earth. Most of the water on Earth is stored for long periods in the oceans, underground, and as ice. Figure $2$ illustrates the average time that an individual water molecule may spend in the Earth’s major water reservoirs. Residence time is a measure of the average time an individual water molecule stays in a particular reservoir. A large amount of the Earth’s water is locked in place in these reservoirs as ice, beneath the ground, and in the ocean, and, thus, is unavailable for short-term cycling (only surface water can evaporate).
There are various processes that occur during the cycling of water, shown in Figure $3$. These processes include the following:
• evaporation/sublimation
• condensation/precipitation
• subsurface water flow
• surface runoff/snowmelt
• streamflow
The water cycle is driven by the sun’s energy as it warms the oceans and other surface waters. This leads to the evaporation (water to water vapor) of liquid surface water and the sublimation (ice to water vapor) of frozen water, which deposits large amounts of water vapor into the atmosphere. Over time, this water vapor condenses into clouds as liquid or frozen droplets and is eventually followed by precipitation (rain or snow), which returns water to the Earth’s surface. Rain eventually permeates into the ground, where it may evaporate again if it is near the surface, flow beneath the surface, or be stored for long periods. More easily observed is surface runoff: the flow of fresh water either from rain or melting ice. Runoff can then make its way through streams and lakes to the oceans or flow directly to the oceans themselves.
Link to Learning
Head to this website to learn more about the world’s fresh water supply.
Rain and surface runoff are major ways in which minerals, including carbon, nitrogen, phosphorus, and sulfur, are cycled from land to water. The environmental effects of runoff will be discussed later as these cycles are described.
The Carbon Cycle
Carbon is the second most abundant element in living organisms. Carbon is present in all organic molecules, and its role in the structure of macromolecules is of primary importance to living organisms. Carbon compounds contain especially high energy, particularly those derived from fossilized organisms, mainly plants, which humans use as fuel. Since the 1800s, the number of countries using massive amounts of fossil fuels has increased. Since the beginning of the Industrial Revolution, global demand for the Earth’s limited fossil fuel supplies has risen; therefore, the amount of carbon dioxide in our atmosphere has increased. This increase in carbon dioxide has been associated with climate change and other disturbances of the Earth’s ecosystems and is a major environmental concern worldwide. Thus, the “carbon footprint” is based on how much carbon dioxide is produced and how much fossil fuel countries consume.
The carbon cycle is most easily studied as two interconnected sub-cycles: one dealing with rapid carbon exchange among living organisms and the other dealing with the long-term cycling of carbon through geologic processes. The entire carbon cycle is shown in Figure $4$.
Link to Learning
Click this link to read information about the United States Carbon Cycle Science Program.
The Biological Carbon Cycle
Living organisms are connected in many ways, even between ecosystems. A good example of this connection is the exchange of carbon between autotrophs and heterotrophs within and between ecosystems by way of atmospheric carbon dioxide. Carbon dioxide is the basic building block that most autotrophs use to build multi-carbon, high energy compounds, such as glucose. The energy harnessed from the sun is used by these organisms to form the covalent bonds that link carbon atoms together. These chemical bonds thereby store this energy for later use in the process of respiration. Most terrestrial autotrophs obtain their carbon dioxide directly from the atmosphere, while marine autotrophs acquire it in the dissolved form (carbonic acid, H2CO3). However carbon dioxide is acquired, a by-product of the process is oxygen. The photosynthetic organisms are responsible for depositing approximately 21 percent oxygen content of the atmosphere that we observe today.
Heterotrophs and autotrophs are partners in biological carbon exchange (especially the primary consumers, largely herbivores). Heterotrophs acquire the high-energy carbon compounds from the autotrophs by consuming them, and breaking them down by respiration to obtain cellular energy, such as ATP. The most efficient type of respiration, aerobic respiration, requires oxygen obtained from the atmosphere or dissolved in water. Thus, there is a constant exchange of oxygen and carbon dioxide between the autotrophs (which need the carbon) and the heterotrophs (which need the oxygen). Gas exchange through the atmosphere and water is one way that the carbon cycle connects all living organisms on Earth.
The Biogeochemical Carbon Cycle
The movement of carbon through the land, water, and air is complex, and in many cases, it occurs much more slowly geologically than as seen between living organisms. Carbon is stored for long periods in what are known as carbon reservoirs, which include the atmosphere, bodies of liquid water (mostly oceans), ocean sediment, soil, land sediments (including fossil fuels), and the Earth’s interior.
As stated, the atmosphere is a major reservoir of carbon in the form of carbon dioxide and is essential to the process of photosynthesis. The level of carbon dioxide in the atmosphere is greatly influenced by the reservoir of carbon in the oceans. The exchange of carbon between the atmosphere and water reservoirs influences how much carbon is found in each location, and each one affects the other reciprocally. Carbon dioxide (CO2) from the atmosphere dissolves in water and combines with water molecules to form carbonic acid, and then it ionizes to carbonate and bicarbonate ions (Figure $5$)
The equilibrium coefficients are such that more than 90 percent of the carbon in the ocean is found as bicarbonate ions. Some of these ions combine with seawater calcium to form calcium carbonate (CaCO3), a major component of marine organism shells. These organisms eventually form sediments on the ocean floor. Over geologic time, the calcium carbonate forms limestone, which comprises the largest carbon reservoir on Earth.
On land, carbon is stored in soil as a result of the decomposition of living organisms (by decomposers) or from weathering of terrestrial rock and minerals. This carbon can be leached into the water reservoirs by surface runoff. Deeper underground, on land and at sea, are fossil fuels: the anaerobically decomposed remains of plants that take millions of years to form. Fossil fuels are considered a non-renewable resource because their use far exceeds their rate of formation. A non-renewable resource, such as fossil fuel, is either regenerated very slowly or not at all. Another way for carbon to enter the atmosphere is from land (including land beneath the surface of the ocean) by the eruption of volcanoes and other geothermal systems. Carbon sediments from the ocean floor are taken deep within the Earth by the process of subduction: the movement of one tectonic plate beneath another. Carbon is released as carbon dioxide when a volcano erupts or from volcanic hydrothermal vents.
Carbon dioxide is also added to the atmosphere by the animal husbandry practices of humans. The large numbers of land animals raised to feed the Earth’s growing population results in increased carbon dioxide levels in the atmosphere due to farming practices and the respiration and methane production. This is another example of how human activity indirectly affects biogeochemical cycles in a significant way. Although much of the debate about the future effects of increasing atmospheric carbon on climate change focuses on fossils fuels, scientists take natural processes, such as volcanoes and respiration, into account as they model and predict the future impact of this increase.
The Nitrogen Cycle
Getting nitrogen into the living world is difficult. Plants and phytoplankton are not equipped to incorporate nitrogen from the atmosphere (which exists as tightly bonded, triple covalent N2) even though this molecule comprises approximately 78 percent of the atmosphere. Nitrogen enters the living world via free-living and symbiotic bacteria, which incorporate nitrogen into their macromolecules through nitrogen fixation (conversion of N2). Cyanobacteria live in most aquatic ecosystems where sunlight is present; they play a key role in nitrogen fixation. Cyanobacteria are able to use inorganic sources of nitrogen to “fix” nitrogen. Rhizobium bacteria live symbiotically in the root nodules of legumes (such as peas, beans, and peanuts) and provide them with the organic nitrogen they need. Free-living bacteria, such as Azotobacter, are also important nitrogen fixers.
Organic nitrogen is especially important to the study of ecosystem dynamics since many ecosystem processes, such as primary production and decomposition, are limited by the available supply of nitrogen. As shown in Figure $6$, the nitrogen that enters living systems by nitrogen fixation is successively converted from organic nitrogen back into nitrogen gas by bacteria. This process occurs in three steps in terrestrial systems: ammonification, nitrification, and denitrification. First, the ammonification process converts nitrogenous waste from living animals or from the remains of dead animals into ammonium (NH4+) by certain bacteria and fungi. Second, the ammonium is converted to nitrites (NO2) by nitrifying bacteria, such as Nitrosomonas, through nitrification. Subsequently, nitrites are converted to nitrates (NO3) by similar organisms. Third, the process of denitrification occurs, whereby bacteria, such as Pseudomonas and Clostridium, convert the nitrates into nitrogen gas, allowing it to re-enter the atmosphere.
Exercise
Which of the following statements about the nitrogen cycle is false?
1. Ammonification converts organic nitrogenous matter from living organisms into ammonium (NH4+).
2. Denitrification by bacteria converts nitrates (NO3) to nitrogen gas (N2).
3. Nitrification by bacteria converts nitrates (NO3) to nitrites (NO2).
4. Nitrogen fixing bacteria convert nitrogen gas (N2) into organic compounds.
Answer
C: Nitrification by bacteria converts nitrates (NO3) to nitrites (NO2).
Human activity can release nitrogen into the environment by two primary means: the combustion of fossil fuels, which releases different nitrogen oxides, and by the use of artificial fertilizers in agriculture, which are then washed into lakes, streams, and rivers by surface runoff. Atmospheric nitrogen is associated with several effects on Earth’s ecosystems including the production of acid rain (as nitric acid, HNO3) and greenhouse gas (as nitrous oxide, N2O) potentially causing climate change. A major effect from fertilizer runoff is saltwater and freshwater eutrophication, a process whereby nutrient runoff causes the excess growth of microorganisms, depleting dissolved oxygen levels and killing ecosystem fauna.
A similar process occurs in the marine nitrogen cycle, where the ammonification, nitrification, and denitrification processes are performed by marine bacteria. Some of this nitrogen falls to the ocean floor as sediment, which can then be moved to land in geologic time by uplift of the Earth’s surface and thereby incorporated into terrestrial rock. Although the movement of nitrogen from rock directly into living systems has been traditionally seen as insignificant compared with nitrogen fixed from the atmosphere, a recent study showed that this process may indeed be significant and should be included in any study of the global nitrogen cycle.1
The Phosphorus Cycle
Phosphorus is an essential nutrient for living processes; it is a major component of nucleic acid and phospholipids, and, as calcium phosphate, makes up the supportive components of our bones. Phosphorus is often the limiting nutrient (necessary for growth) in aquatic ecosystems (Figure $7$).
Phosphorus occurs in nature as the phosphate ion (PO43−). In addition to phosphate runoff as a result of human activity, natural surface runoff occurs when it is leached from phosphate-containing rock by weathering, thus sending phosphates into rivers, lakes, and the ocean. This rock has its origins in the ocean. Phosphate-containing ocean sediments form primarily from the bodies of ocean organisms and from their excretions. However, in remote regions, volcanic ash, aerosols, and mineral dust may also be significant phosphate sources. This sediment then is moved to land over geologic time by the uplifting of areas of the Earth’s surface.
Phosphorus is also reciprocally exchanged between phosphate dissolved in the ocean and marine ecosystems. The movement of phosphate from the ocean to the land and through the soil is extremely slow, with the average phosphate ion having an oceanic residence time between 20,000 and 100,000 years.
Excess phosphorus and nitrogen that enters these ecosystems from fertilizer runoff and from sewage causes excessive growth of microorganisms and depletes the dissolved oxygen, which leads to the death of many ecosystem fauna, such as shellfish and finfish. This process is responsible for dead zones in lakes and at the mouths of many major rivers (Figure $8$).
A dead zone is an area within a freshwater or marine ecosystem where large areas are depleted of their normal flora and fauna; these zones can be caused by eutrophication, oil spills, dumping of toxic chemicals, and other human activities. The number of dead zones has been increasing for several years, and more than 400 of these zones were present as of 2008. One of the worst dead zones is off the coast of the United States in the Gulf of Mexico, where fertilizer runoff from the Mississippi River basin has created a dead zone of over 8463 square miles. Phosphate and nitrate runoff from fertilizers also negatively affect several lake and bay ecosystems including the Chesapeake Bay in the eastern United States.
Everyday Connection: Chesapeake Bay
The Chesapeake Bay has long been valued as one of the most scenic areas on Earth; it is now in distress and is recognized as a declining ecosystem. In the 1970s, the Chesapeake Bay was one of the first ecosystems to have identified dead zones, which continue to kill many fish and bottom-dwelling species, such as clams, oysters, and worms. Several species have declined in the Chesapeake Bay due to surface water runoff containing excess nutrients from artificial fertilizer used on land. The source of the fertilizers (with high nitrogen and phosphate content) is not limited to agricultural practices. There are many nearby urban areas and more than 150 rivers and streams empty into the bay that are carrying fertilizer runoff from lawns and gardens. Thus, the decline of the Chesapeake Bay is a complex issue and requires the cooperation of industry, agriculture, and everyday homeowners.
Of particular interest to conservationists is the oyster population; it is estimated that more than 200,000 acres of oyster reefs existed in the bay in the 1700s, but that number has now declined to only 36,000 acres. Oyster harvesting was once a major industry for Chesapeake Bay, but it declined 88 percent between 1982 and 2007. This decline was due not only to fertilizer runoff and dead zones but also to overharvesting. Oysters require a certain minimum population density because they must be in close proximity to reproduce. Human activity has altered the oyster population and locations, greatly disrupting the ecosystem.
The restoration of the oyster population in the Chesapeake Bay has been ongoing for several years with mixed success. Not only do many people find oysters good to eat, but they also clean up the bay. Oysters are filter feeders, and as they eat, they clean the water around them. In the 1700s, it was estimated that it took only a few days for the oyster population to filter the entire volume of the bay. Today, with changed water conditions, it is estimated that the present population would take nearly a year to do the same job.
Restoration efforts have been ongoing for several years by non-profit organizations, such as the Chesapeake Bay Foundation. The restoration goal is to find a way to increase population density so the oysters can reproduce more efficiently. Many disease-resistant varieties (developed at the Virginia Institute of Marine Science for the College of William and Mary) are now available and have been used in the construction of experimental oyster reefs. Efforts to clean and restore the bay by Virginia and Delaware have been hampered because much of the pollution entering the bay comes from other states, which stresses the need for inter-state cooperation to gain successful restoration.
The new, hearty oyster strains have also spawned a new and economically viable industry—oyster aquaculture—which not only supplies oysters for food and profit, but also has the added benefit of cleaning the bay.
The Sulfur Cycle
Sulfur is an essential element for the macromolecules of living things. As a part of the amino acid cysteine, it is involved in the formation of disulfide bonds within proteins, which help to determine their 3-D folding patterns, and hence their functions. As shown in Figure $10$, sulfur cycles between the oceans, land, and atmosphere. Atmospheric sulfur is found in the form of sulfur dioxide (SO2) and enters the atmosphere in three ways: from the decomposition of organic molecules, from volcanic activity and geothermal vents, and from the burning of fossil fuels by humans.
On land, sulfur is deposited in four major ways: precipitation, direct fallout from the atmosphere, rock weathering, and geothermal vents (Figure $11$). Atmospheric sulfur is found in the form of sulfur dioxide (SO2), and as rain falls through the atmosphere, sulfur is dissolved in the form of weak sulfuric acid (H2SO4). Sulfur can also fall directly from the atmosphere in a process called fallout. Also, the weathering of sulfur-containing rocks releases sulfur into the soil. These rocks originate from ocean sediments that are moved to land by the geologic uplifting of ocean sediments. Terrestrial ecosystems can then make use of these soil sulfates ($\text{SO}_4^{2-}$), and upon the death and decomposition of these organisms, release the sulfur back into the atmosphere as hydrogen sulfide (H2S) gas.
Sulfur enters the ocean via runoff from land, from atmospheric fallout, and from underwater geothermal vents. Some ecosystems rely on chemoautotrophs using sulfur as a biological energy source. This sulfur then supports marine ecosystems in the form of sulfates.
Human activities have played a major role in altering the balance of the global sulfur cycle. The burning of large quantities of fossil fuels, especially from coal, releases larger amounts of hydrogen sulfide gas into the atmosphere. As rain falls through this gas, it creates the phenomenon known as acid rain. Acid rain is corrosive rain caused by rainwater falling to the ground through sulfur dioxide gas, turning it into weak sulfuric acid, which causes damage to aquatic ecosystems. Acid rain damages the natural environment by lowering the pH of lakes, which kills many of the resident fauna; it also affects the man-made environment through the chemical degradation of buildings. For example, many marble monuments, such as the Lincoln Memorial in Washington, DC, have suffered significant damage from acid rain over the years. These examples show the wide-ranging effects of human activities on our environment and the challenges that remain for our future.
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Click this link to learn more about global climate change.
Summary
Mineral nutrients are cycled through ecosystems and their environment. Of particular importance are water, carbon, nitrogen, phosphorus, and sulfur. All of these cycles have major impacts on ecosystem structure and function. As human activities have caused major disturbances to these cycles, their study and modeling is especially important. A variety of human activities, such as pollution, oil spills, and other events have damaged ecosystems, potentially causing global climate change. The health of Earth depends on understanding these cycles and how to protect the environment from irreversible damage.
Footnotes
1. 1 Scott L. Morford, Benjamin Z. Houlton, and Randy A. Dahlgren, “Increased Forest Ecosystem Carbon and Nitrogen Storage from Nitrogen Rich Bedrock,” Nature 477, no. 7362 (2011): 78–81.
Glossary
acid rain
corrosive rain caused by rainwater falling to the ground through sulfur dioxide gas, turning it into weak sulfuric acid; can damage structures and ecosystems
biogeochemical cycle
cycling of mineral nutrients through ecosystems and through the non-living world
dead zone
area within an ecosystem in lakes and near the mouths of rivers where large areas of ecosystems are depleted of their normal flora and fauna; these zones can be caused by eutrophication, oil spills, dumping of toxic chemicals, and other human activities
eutrophication
process whereby nutrient runoff causes the excess growth of microorganisms, depleting dissolved oxygen levels and killing ecosystem fauna
fallout
direct deposit of solid minerals on land or in the ocean from the atmosphere
hydrosphere
area of the Earth where water movement and storage occurs
non-renewable resource
resource, such as fossil fuel, that is either regenerated very slowly or not at all
residence time
measure of the average time an individual water molecule stays in a particular reservoir
subduction
movement of one tectonic plate beneath another | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/56%3A_Dynamics_of_Ecosystems/56.01%3A_Biogeochemical_Cycles.txt |
Learning Objectives
• Describe how organisms acquire energy in a food web and in associated food chains
• Explain how the efficiency of energy transfers between trophic levels affects ecosystem structure and dynamics
• Discuss trophic levels and how ecological pyramids are used to model them
All living things require energy in one form or another. Energy is required by most complex metabolic pathways (often in the form of adenosine triphosphate, ATP), especially those responsible for building large molecules from smaller compounds, and life itself is an energy-driven process. Living organisms would not be able to assemble macromolecules (proteins, lipids, nucleic acids, and complex carbohydrates) from their monomeric subunits without a constant energy input.
It is important to understand how organisms acquire energy and how that energy is passed from one organism to another through food webs and their constituent food chains. Food webs illustrate how energy flows directionally through ecosystems, including how efficiently organisms acquire it, use it, and how much remains for use by other organisms of the food web.
How Organisms Acquire Energy in a Food Web
Energy is acquired by living things in three ways: photosynthesis, chemosynthesis, and the consumption and digestion of other living or previously living organisms by heterotrophs.
Photosynthetic and chemosynthetic organisms are both grouped into a category known as autotrophs: organisms capable of synthesizing their own food (more specifically, capable of using inorganic carbon as a carbon source). Photosynthetic autotrophs (photoautotrophs) use sunlight as an energy source, whereas chemosynthetic autotrophs (chemoautotrophs) use inorganic molecules as an energy source. Autotrophs are critical for all ecosystems. Without these organisms, energy would not be available to other living organisms and life itself would not be possible.
Photoautotrophs, such as plants, algae, and photosynthetic bacteria, serve as the energy source for a majority of the world’s ecosystems. These ecosystems are often described by grazing food webs. Photoautotrophs harness the solar energy of the sun by converting it to chemical energy in the form of ATP (and NADP). The energy stored in ATP is used to synthesize complex organic molecules, such as glucose.
Chemoautotrophs are primarily bacteria that are found in rare ecosystems where sunlight is not available, such as in those associated with dark caves or hydrothermal vents at the bottom of the ocean (Figure $1$). Many chemoautotrophs in hydrothermal vents use hydrogen sulfide (H2S), which is released from the vents as a source of chemical energy. This allows chemoautotrophs to synthesize complex organic molecules, such as glucose, for their own energy and in turn supplies energy to the rest of the ecosystem.
Productivity within Trophic Levels
Productivity within an ecosystem can be defined as the percentage of energy entering the ecosystem incorporated into biomass in a particular trophic level. Biomass is the total mass, in a unit area at the time of measurement, of living or previously living organisms within a trophic level. Ecosystems have characteristic amounts of biomass at each trophic level. For example, in the English Channel ecosystem the primary producers account for a biomass of 4 g/m2 (grams per meter squared), while the primary consumers exhibit a biomass of 21 g/m2.
The productivity of the primary producers is especially important in any ecosystem because these organisms bring energy to other living organisms by photoautotrophy or chemoautotrophy. The rate at which photosynthetic primary producers incorporate energy from the sun is called gross primary productivity. An example of gross primary productivity is shown in the compartment diagram of energy flow within the Silver Springs aquatic ecosystem as shown (Figure 46.1.7). In this ecosystem, the total energy accumulated by the primary producers (gross primary productivity) was shown to be 20,810 kcal/m2/yr.
Because all organisms need to use some of this energy for their own functions (like respiration and resulting metabolic heat loss) scientists often refer to the net primary productivity of an ecosystem. Net primary productivity is the energy that remains in the primary producers after accounting for the organisms’ respiration and heat loss. The net productivity is then available to the primary consumers at the next trophic level. In our Silver Spring example, 13,187 of the 20,810 kcal/m2/yr were used for respiration or were lost as heat, leaving 7,632 kcal/m2/yr of energy for use by the primary consumers.
Ecological Efficiency: The Transfer of Energy between Trophic Levels
As illustrated in Figure 46.1.7, large amounts of energy are lost from the ecosystem from one trophic level to the next level as energy flows from the primary producers through the various trophic levels of consumers and decomposers. The main reason for this loss is the second law of thermodynamics, which states that whenever energy is converted from one form to another, there is a tendency toward disorder (entropy) in the system. In biologic systems, this means a great deal of energy is lost as metabolic heat when the organisms from one trophic level consume the next level. In the Silver Springs ecosystem example (Figure 46.1.7), we see that the primary consumers produced 1103 kcal/m2/yr from the 7618 kcal/m2/yr of energy available to them from the primary producers. The measurement of energy transfer efficiency between two successive trophic levels is termed the trophic level transfer efficiency (TLTE) and is defined by the formula:
$\text{TLTE} = \frac{\text{production at present trophic level}}{\text{production at previous trophic level}} * 100 \nonumber$
In Silver Springs, the TLTE between the first two trophic levels was approximately 14.8 percent. The low efficiency of energy transfer between trophic levels is usually the major factor that limits the length of food chains observed in a food web. The fact is, after four to six energy transfers, there is not enough energy left to support another trophic level. In the Lake Ontario example shown in Figure 46.1.5, only three energy transfers occurred between the primary producer, (green algae), and the apex consumer (Chinook salmon).
Ecologists have many different methods of measuring energy transfers within ecosystems. Some transfers are easier or more difficult to measure depending on the complexity of the ecosystem and how much access scientists have to observe the ecosystem. In other words, some ecosystems are more difficult to study than others, and sometimes the quantification of energy transfers has to be estimated.
Another main parameter that is important in characterizing energy flow within an ecosystem is the net production efficiency. Net production efficiency (NPE) allows ecologists to quantify how efficiently organisms of a particular trophic level incorporate the energy they receive into biomass; it is calculated using the following formula:
$\text{NPE} = \frac{\text{net consumer productivity}}{\text{assimilation}} * 100 \nonumber$
Net consumer productivity is the energy content available to the organisms of the next trophic level. Assimilation is the biomass (energy content generated per unit area) of the present trophic level after accounting for the energy lost due to incomplete ingestion of food, energy used for respiration, and energy lost as waste. Incomplete ingestion refers to the fact that some consumers eat only a part of their food. For example, when a lion kills an antelope, it will eat everything except the hide and bones. The lion is missing the energy-rich bone marrow inside the bone, so the lion does not make use of all the calories its prey could provide.
Thus, NPE measures how efficiently each trophic level uses and incorporates the energy from its food into biomass to fuel the next trophic level. In general, cold-blooded animals (ectotherms), such as invertebrates, fish, amphibians, and reptiles, use less of the energy they obtain for respiration and heat than warm-blooded animals (endotherms), such as birds and mammals. The extra heat generated in endotherms, although an advantage in terms of the activity of these organisms in colder environments, is a major disadvantage in terms of NPE. Therefore, many endotherms have to eat more often than ectotherms to get the energy they need for survival. In general, NPE for ectotherms is an order of magnitude (10x) higher than for endotherms. For example, the NPE for a caterpillar eating leaves has been measured at 18 percent, whereas the NPE for a squirrel eating acorns may be as low as 1.6 percent.
The inefficiency of energy use by warm-blooded animals has broad implications for the world's food supply. It is widely accepted that the meat industry uses large amounts of crops to feed livestock, and because the NPE is low, much of the energy from animal feed is lost. For example, it costs about 1¢ to produce 1000 dietary calories (kcal) of corn or soybeans, but approximately $0.19 to produce a similar number of calories growing cattle for beef consumption. The same energy content of milk from cattle is also costly, at approximately$0.16 per 1000 kcal. Much of this difference is due to the low NPE of cattle. Thus, there has been a growing movement worldwide to promote the consumption of non-meat and non-dairy foods so that less energy is wasted feeding animals for the meat industry.
Modeling Ecosystems Energy Flow: Ecological Pyramids
The structure of ecosystems can be visualized with ecological pyramids, which were first described by the pioneering studies of Charles Elton in the 1920s. Ecological pyramids show the relative amounts of various parameters (such as number of organisms, energy, and biomass) across trophic levels.
Pyramids of numbers can be either upright or inverted, depending on the ecosystem. As shown in Figure $2$, typical grassland during the summer has a base of many plants and the numbers of organisms decrease at each trophic level. However, during the summer in a temperate forest, the base of the pyramid consists of few trees compared with the number of primary consumers, mostly insects. Because trees are large, they have great photosynthetic capability, and dominate other plants in this ecosystem to obtain sunlight. Even in smaller numbers, primary producers in forests are still capable of supporting other trophic levels.
Another way to visualize ecosystem structure is with pyramids of biomass. This pyramid measures the amount of energy converted into living tissue at the different trophic levels. Using the Silver Springs ecosystem example, this data exhibits an upright biomass pyramid (Figure $2$), whereas the pyramid from the English Channel example is inverted. The plants (primary producers) of the Silver Springs ecosystem make up a large percentage of the biomass found there. However, the phytoplankton in the English Channel example make up less biomass than the primary consumers, the zooplankton. As with inverted pyramids of numbers, this inverted pyramid is not due to a lack of productivity from the primary producers, but results from the high turnover rate of the phytoplankton. The phytoplankton are consumed rapidly by the primary consumers, thus, minimizing their biomass at any particular point in time. However, phytoplankton reproduce quickly, thus they are able to support the rest of the ecosystem.
Pyramid ecosystem modeling can also be used to show energy flow through the trophic levels. Notice that these numbers are the same as those used in the energy flow compartment diagram in Figure 46.1.7. Pyramids of energy are always upright, and an ecosystem without sufficient primary productivity cannot be supported. All types of ecological pyramids are useful for characterizing ecosystem structure. However, in the study of energy flow through the ecosystem, pyramids of energy are the most consistent and representative models of ecosystem structure (Figure $2$).
Exercise
Pyramids depicting the number of organisms or biomass may be inverted, upright, or even diamond-shaped. Energy pyramids, however, are always upright. Why?
Answer
Pyramids of organisms may be inverted or diamond-shaped because a large organism, such as a tree, can sustain many smaller organisms. Likewise, a low biomass of organisms can sustain a larger biomass at the next trophic level because the organisms reproduce rapidly and thus supply continuous nourishment. Energy pyramids, however, must always be upright because of the laws of thermodynamics. The first law of thermodynamics states that energy can neither be created nor destroyed; thus, each trophic level must acquire energy from the trophic level below. The second law of thermodynamics states that, during the transfer of energy, some energy is always lost as heat; thus, less energy is available at each higher trophic level.
Consequences of Food Webs: Biological Magnification
One of the most important environmental consequences of ecosystem dynamics is biomagnification. Biomagnification is the increasing concentration of persistent, toxic substances in organisms at each trophic level, from the primary producers to the apex consumers. Many substances have been shown to bioaccumulate, including classical studies with the pesticide dichlorodiphenyltrichloroethane (DDT), which was published in the 1960s bestseller, Silent Spring, by Rachel Carson. DDT was a commonly used pesticide before its dangers became known. In some aquatic ecosystems, organisms from each trophic level consumed many organisms of the lower level, which caused DDT to increase in birds (apex consumers) that ate fish. Thus, the birds accumulated sufficient amounts of DDT to cause fragility in their eggshells. This effect increased egg breakage during nesting and was shown to have adverse effects on these bird populations. The use of DDT was banned in the United States in the 1970s.
Other substances that biomagnify are polychlorinated biphenyls (PCBs), which were used in coolant liquids in the United States until their use was banned in 1979, and heavy metals, such as mercury, lead, and cadmium. These substances were best studied in aquatic ecosystems, where fish species at different trophic levels accumulate toxic substances brought through the ecosystem by the primary producers. As illustrated in a study performed by the National Oceanic and Atmospheric Administration (NOAA) in the Saginaw Bay of Lake Huron (Figure $3$), PCB concentrations increased from the ecosystem’s primary producers (phytoplankton) through the different trophic levels of fish species. The apex consumer (walleye) has more than four times the amount of PCBs compared to phytoplankton. Also, based on results from other studies, birds that eat these fish may have PCB levels at least one order of magnitude higher than those found in the lake fish.
Other concerns have been raised by the accumulation of heavy metals, such as mercury and cadmium, in certain types of seafood. The United States Environmental Protection Agency (EPA) recommends that pregnant women and young children should not consume any swordfish, shark, king mackerel, or tilefish because of their high mercury content. These individuals are advised to eat fish low in mercury: salmon, tilapia, shrimp, pollock, and catfish. Biomagnification is a good example of how ecosystem dynamics can affect our everyday lives, even influencing the food we eat.
Summary
Organisms in an ecosystem acquire energy in a variety of ways, which is transferred between trophic levels as the energy flows from the bottom to the top of the food web, with energy being lost at each transfer. The efficiency of these transfers is important for understanding the different behaviors and eating habits of warm-blooded versus cold-blooded animals. Modeling of ecosystem energy is best done with ecological pyramids of energy, although other ecological pyramids provide other vital information about ecosystem structure.
Glossary
assimilation
biomass consumed and assimilated from the previous trophic level after accounting for the energy lost due to incomplete ingestion of food, energy used for respiration, and energy lost as waste
biomagnification
increasing concentrations of persistent, toxic substances in organisms at each trophic level, from the primary producers to the apex consumers
biomass
total weight, at the time of measurement, of living or previously living organisms in a unit area within a trophic level
chemoautotroph
organism capable of synthesizing its own food using energy from inorganic molecules
ecological pyramid
(also, Eltonian pyramid) graphical representation of different trophic levels in an ecosystem based of organism numbers, biomass, or energy content
gross primary productivity
rate at which photosynthetic primary producers incorporate energy from the sun
net consumer productivity
energy content available to the organisms of the next trophic level
net primary productivity
energy that remains in the primary producers after accounting for the organisms’ respiration and heat loss
net production efficiency (NPE)
measure of the ability of a trophic level to convert the energy it receives from the previous trophic level into biomass
trophic level transfer efficiency (TLTE)
energy transfer efficiency between two successive trophic levels | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/56%3A_Dynamics_of_Ecosystems/56.02%3A_The_Flow_of_Energy_in_Ecosystems.txt |
• 57.1: Ecosystem Effects of Sun, Wind and Water
• 57.2: Earth's Biomes
The Earth’s biomes are categorized into two major groups: terrestrial and aquatic. Terrestrial biomes are based on land, while aquatic biomes include both ocean and freshwater biomes. The eight major terrestrial biomes on Earth are each distinguished by characteristic temperatures and amount of precipitation. Comparing the annual totals of precipitation and fluctuations in precipitation from one biome to another provides clues as to the importance of abiotic factors in the distribution of biomes
• 57.3: Freshwater Habitats
• 57.4: Marine Habitats
Like terrestrial biomes, aquatic biomes are influenced by a series of abiotic factors. The aquatic medium—water— has different physical and chemical properties than air, however. Even if the water in a pond or other body of water is perfectly clear (there are no suspended particles), water, on its own, absorbs light. As one descends into a deep body of water, there will eventually be a depth which the sunlight cannot reach.
• 57.5: Human Impacts of the Biosphere- Pollution and Resource Depletion
All biomes are universally affected by global conditions, such as climate, that ultimately shape each biome’s environment. Scientists who study climate have noted a series of marked changes that have gradually become increasingly evident during the last sixty years. Global climate change is the term used to describe altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide.
• 57.6: Human Impacts on the Biosphere- Climate Change
57: The Biosphere and Human Impacts
Skills to Develop
• Identify the two major abiotic factors that determine terrestrial biomes
• Recognize distinguishing characteristics of each of the eight major terrestrial biomes
The Earth’s biomes are categorized into two major groups: terrestrial and aquatic. Terrestrial biomes are based on land, while aquatic biomes include both ocean and freshwater biomes. The eight major terrestrial biomes on Earth are each distinguished by characteristic temperatures and amount of precipitation. Comparing the annual totals of precipitation and fluctuations in precipitation from one biome to another provides clues as to the importance of abiotic factors in the distribution of biomes. Temperature variation on a daily and seasonal basis is also important for predicting the geographic distribution of the biome and the vegetation type in the biome. The distribution of these biomes shows that the same biome can occur in geographically distinct areas with similar climates (Figure \(1\)).
Art Connection
Which of the following statements about biomes is false?
1. Chaparral is dominated by shrubs.
2. Savannas and temperate grasslands are dominated by grasses.
3. Boreal forests are dominated by deciduous trees.
4. Lichens are common in the arctic tundra.
Tropical Wet Forest
Tropical wet forests are also referred to as tropical rainforests. This biome is found in equatorial regions (Figure \(1\)). The vegetation is characterized by plants with broad leaves that fall off throughout the year. Unlike the trees of deciduous forests, the trees in this biome do not have a seasonal loss of leaves associated with variations in temperature and sunlight; these forests are “evergreen” year-round.
The temperature and sunlight profiles of tropical wet forests are very stable in comparison to that of other terrestrial biomes, with the temperatures ranging from 20 °C to 34 °C (68 °F to 93 °F). When one compares the annual temperature variation of tropical wet forests with that of other forest biomes, the lack of seasonal temperature variation in the tropical wet forest becomes apparent. This lack of seasonality leads to year-round plant growth, rather than the seasonal (spring, summer, and fall) growth seen in other biomes. In contrast to other ecosystems, tropical ecosystems do not have long days and short days during the yearly cycle. Instead, a constant daily amount of sunlight (11–12 hrs per day) provides more solar radiation, thereby, a longer period of time for plant growth.
The annual rainfall in tropical wet forests ranges from 125 to 660 cm (50–200 in) with some monthly variation. While sunlight and temperature remain fairly consistent, annual rainfall is highly variable. Tropical wet forests have wet months in which there can be more than 30 cm (11–12 in) of precipitation, as well as dry months in which there are fewer than 10 cm (3.5 in) of rainfall. However, the driest month of a tropical wet forest still exceeds the annual rainfall of some other biomes, such as deserts.
Tropical wet forests have high net primary productivity because the annual temperatures and precipitation values in these areas are ideal for plant growth. Therefore, the extensive biomass present in the tropical wet forest leads to plant communities with very high species diversities (Figure \(2\)). Tropical wet forests have more species of trees than any other biome; on average between 100 and 300 species of trees are present in a single hectare (2.5 acres) of South America. One way to visualize this is to compare the distinctive horizontal layers within the tropical wet forest biome. On the forest floor is a sparse layer of plants and decaying plant matter. Above that is an understory of short shrubby foliage. A layer of trees rises above this understory and is topped by a closed upper canopy—the uppermost overhead layer of branches and leaves. Some additional trees emerge through this closed upper canopy. These layers provide diverse and complex habitats for the variety of plants, fungi, animals, and other organisms within the tropical wet forests. For instance, epiphytes are plants that grow on other plants, which typically are not harmed. Epiphytes are found throughout tropical wet forest biomes. Many species of animals use the variety of plants and the complex structure of the tropical wet forests for food and shelter. Some organisms live several meters above ground and have adapted to this arboreal lifestyle.
Savannas
Savannas are grasslands with scattered trees, and they are located in Africa, South America, and northern Australia (Figure \(3\)). Savannas are hot, tropical areas with temperatures averaging from 24 °C to 29 °C (75 °F to 84 °F) and an annual rainfall of 10–40 cm (3.9–15.7 in). Savannas have an extensive dry season; for this reason, forest trees do not grow as well as they do in the tropical wet forest (or other forest biomes). As a result, within the grasses and forbs (herbaceous flowering plants) that dominate the savanna, there are relatively few trees (Figure \(3\)). Since fire is an important source of disturbance in this biome, plants have evolved well-developed root systems that allow them to quickly re-sprout after a fire.
Subtropical Deserts
Subtropical deserts exist between 15 ° and 30 ° north and south latitude and are centered on the Tropics of Cancer and Capricorn (Figure \(4\)). This biome is very dry; in some years, evaporation exceeds precipitation. Subtropical hot deserts can have daytime soil surface temperatures above 60 °C (140 °F) and nighttime temperatures approaching 0 °C (32 °F). In cold deserts, temperatures can be as high as 25 °C and can drop below -30 °C (-22 °F). Subtropical deserts are characterized by low annual precipitation of fewer than 30 cm (12 in) with little monthly variation and lack of predictability in rainfall. In some cases, the annual rainfall can be as low as 2 cm (0.8 in) in subtropical deserts located in central Australia (“the Outback”) and northern Africa.
The vegetation and low animal diversity of this biome is closely related to this low and unpredictable precipitation. Very dry deserts lack perennial vegetation that lives from one year to the next; instead, many plants are annuals that grow quickly and reproduce when rainfall does occur, then they die. Many other plants in these areas are characterized by having a number of adaptations that conserve water, such as deep roots, reduced foliage, and water-storing stems (Figure \(4\)). Seed plants in the desert produce seeds that can be in dormancy for extended periods between rains. Adaptations in desert animals include nocturnal behavior and burrowing.
Chaparral
The chaparral is also called the scrub forest and is found in California, along the Mediterranean Sea, and along the southern coast of Australia (Figure \(5\)). The annual rainfall in this biome ranges from 65 cm to 75 cm (25.6–29.5 in), and the majority of the rain falls in the winter. Summers are very dry and many chaparral plants are dormant during the summertime. The chaparral vegetation, shown in Figure \(5\), is dominated by shrubs and is adapted to periodic fires, with some plants producing seeds that only germinate after a hot fire. The ashes left behind after a fire are rich in nutrients like nitrogen that fertilize the soil and promote plant regrowth.
Temperate Grasslands
Temperate grasslands are found throughout central North America, where they are also known as prairies; they are also in Eurasia, where they are known as steppes (Figure \(6\)). Temperate grasslands have pronounced annual fluctuations in temperature with hot summers and cold winters. The annual temperature variation produces specific growing seasons for plants. Plant growth is possible when temperatures are warm enough to sustain plant growth and when ample water is available, which occurs in the spring, summer, and fall. During much of the winter, temperatures are low, and water, which is stored in the form of ice, is not available for plant growth.
Annual precipitation ranges from 25 cm to 75 cm (9.8–29.5 in). Because of relatively lower annual precipitation in temperate grasslands, there are few trees except for those found growing along rivers or streams. The dominant vegetation tends to consist of grasses and some prairies sustain populations of grazing animals Figure \(6\). The vegetation is very dense and the soils are fertile because the subsurface of the soil is packed with the roots and rhizomes (underground stems) of these grasses. The roots and rhizomes act to anchor plants into the ground and replenish the organic material (humus) in the soil when they die and decay.
Fires, mainly caused by lightning, are a natural disturbance in temperate grasslands. When fire is suppressed in temperate grasslands, the vegetation eventually converts to scrub and dense forests. Often, the restoration or management of temperate grasslands requires the use of controlled burns to suppress the growth of trees and maintain the grasses.
Temperate Forests
Temperate forests are the most common biome in eastern North America, Western Europe, Eastern Asia, Chile, and New Zealand (Figure \(7\)). This biome is found throughout mid-latitude regions. Temperatures range between -30 °C and 30 °C (-22 °F to 86 °F) and drop to below freezing on an annual basis. These temperatures mean that temperate forests have defined growing seasons during the spring, summer, and early fall. Precipitation is relatively constant throughout the year and ranges between 75 cm and 150 cm (29.5–59 in).
Because of the moderate annual rainfall and temperatures, deciduous trees are the dominant plant in this biome (Figure \(7\)). Deciduous trees lose their leaves each fall and remain leafless in the winter. Thus, no photosynthesis occurs in the deciduous trees during the dormant winter period. Each spring, new leaves appear as the temperature increases. Because of the dormant period, the net primary productivity of temperate forests is less than that of tropical wet forests. In addition, temperate forests show less diversity of tree species than tropical wet forest biomes.
The trees of the temperate forests leaf out and shade much of the ground; however, this biome is more open than tropical wet forests because trees in the temperate forests do not grow as tall as the trees in tropical wet forests. The soils of the temperate forests are rich in inorganic and organic nutrients. This is due to the thick layer of leaf litter on forest floors. As this leaf litter decays, nutrients are returned to the soil. The leaf litter also protects soil from erosion, insulates the ground, and provides habitats for invertebrates (such as the pill bug or roly-poly, Armadillidium vulgare) and their predators, such as the red-backed salamander (Plethodon cinereus).
Boreal Forests
The boreal forest, also known as taiga or coniferous forest, is found south of the Arctic Circle and across most of Canada, Alaska, Russia, and northern Europe (Figure \(8\)). This biome has cold, dry winters and short, cool, wet summers. The annual precipitation is from 40 cm to 100 cm (15.7–39 in) and usually takes the form of snow. Little evaporation occurs because of the cold temperatures.
The long and cold winters in the boreal forest have led to the predominance of cold-tolerant cone-bearing plants. These are evergreen coniferous trees like pines, spruce, and fir, which retain their needle-shaped leaves year-round. Evergreen trees can photosynthesize earlier in the spring than deciduous trees because less energy from the sun is required to warm a needle-like leaf than a broad leaf. This benefits evergreen trees, which grow faster than deciduous trees in the boreal forest. In addition, soils in boreal forest regions tend to be acidic with little available nitrogen. Leaves are a nitrogen-rich structure and deciduous trees must produce a new set of these nitrogen-rich structures each year. Therefore, coniferous trees that retain nitrogen-rich needles may have a competitive advantage over the broad-leafed deciduous trees.
The net primary productivity of boreal forests is lower than that of temperate forests and tropical wet forests. The aboveground biomass of boreal forests is high because these slow-growing tree species are long lived and accumulate standing biomass over time. Plant species diversity is less than that seen in temperate forests and tropical wet forests. Boreal forests lack the pronounced elements of the layered forest structure seen in tropical wet forests. The structure of a boreal forest is often only a tree layer and a ground layer (Figure \(8\)). When conifer needles are dropped, they decompose more slowly than broad leaves; therefore, fewer nutrients are returned to the soil to fuel plant growth.
Arctic Tundra
The Arctic tundra lies north of the subarctic boreal forest and is located throughout the Arctic regions of the northern hemisphere (Figure \(9\)). The average winter temperature is -34 °C (-34 °F) and the average summer temperature is from 3 °C to 12 °C (37 °F–52 °F). Plants in the arctic tundra have a very short growing season of approximately 10–12 weeks. However, during this time, there are almost 24 hours of daylight and plant growth is rapid. The annual precipitation of the Arctic tundra is very low with little annual variation in precipitation. And, as in the boreal forests, there is little evaporation due to the cold temperatures.
Plants in the Arctic tundra are generally low to the ground (Figure \(9\)). There is little species diversity, low net primary productivity, and low aboveground biomass. The soils of the Arctic tundra may remain in a perennially frozen state referred to as permafrost. The permafrost makes it impossible for roots to penetrate deep into the soil and slows the decay of organic matter, which inhibits the release of nutrients from organic matter. During the growing season, the ground of the Arctic tundra can be completely covered with plants or lichens.
Link to Learning
Watch this Assignment Discovery: Biomes video for an overview of biomes. To explore further, select one of the biomes on the extended playlist: desert, savanna, temperate forest, temperate grassland, tropic, tundra.
Summary
The Earth has terrestrial biomes and aquatic biomes. Aquatic biomes include both freshwater and marine environments. There are eight major terrestrial biomes: tropical wet forests, savannas, subtropical deserts, chaparral, temperate grasslands, temperate forests, boreal forests, and Arctic tundra. The same biome can occur in different geographic locations with similar climates. Temperature and precipitation, and variations in both, are key abiotic factors that shape the composition of animal and plant communities in terrestrial biomes. Some biomes, such as temperate grasslands and temperate forests, have distinct seasons, with cold weather and hot weather alternating throughout the year. In warm, moist biomes, such as the tropical wet forest, net primary productivity is high, as warm temperatures, abundant water, and a year-round growing season fuel plant growth. Other biomes, such as deserts and tundra, have low primary productivity due to extreme temperatures and a shortage of available water.
Art Connections
Figure \(1\): Which of the following statements about biomes is false?
1. Chaparral is dominated by shrubs.
2. Savannas and temperate grasslands are dominated by grasses.
3. Boreal forests are dominated by deciduous trees.
4. Lichens are common in the arctic tundra.
Answer
C. Boreal forests are not dominated by deciduous trees.
Glossary
canopy
branches and foliage of trees that form a layer of overhead coverage in a forest
permafrost
perennially frozen portion of the Arctic tundra soil | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.02%3A_Earth%27s_Biomes.txt |
Skills to Develop
• Describe the effects of abiotic factors on the composition of plant and animal communities in aquatic biomes
• Compare and contrast the characteristics of the ocean zones
• Summarize the characteristics of standing water and flowing water freshwater biomes
Abiotic Factors Influencing Aquatic Biomes
Like terrestrial biomes, aquatic biomes are influenced by a series of abiotic factors. The aquatic medium—water— has different physical and chemical properties than air, however. Even if the water in a pond or other body of water is perfectly clear (there are no suspended particles), water, on its own, absorbs light. As one descends into a deep body of water, there will eventually be a depth which the sunlight cannot reach. While there are some abiotic and biotic factors in a terrestrial ecosystem that might obscure light (like fog, dust, or insect swarms), usually these are not permanent features of the environment. The importance of light in aquatic biomes is central to the communities of organisms found in both freshwater and marine ecosystems. In freshwater systems, stratification due to differences in density is perhaps the most critical abiotic factor and is related to the energy aspects of light. The thermal properties of water (rates of heating and cooling) are significant to the function of marine systems and have major impacts on global climate and weather patterns. Marine systems are also influenced by large-scale physical water movements, such as currents; these are less important in most freshwater lakes.
The ocean is categorized by several areas or zones (Figure \(1\)). All of the ocean’s open water is referred to as the pelagic realm (or zone). The benthic realm (or zone) extends along the ocean bottom from the shoreline to the deepest parts of the ocean floor. Within the pelagic realm is the photic zone, which is the portion of the ocean that light can penetrate (approximately 200 m or 650 ft). At depths greater than 200 m, light cannot penetrate; thus, this is referred to as the aphotic zone. The majority of the ocean is aphotic and lacks sufficient light for photosynthesis. The deepest part of the ocean, the Challenger Deep (in the Mariana Trench, located in the western Pacific Ocean), is about 11,000 m (about 6.8 mi) deep. To give some perspective on the depth of this trench, the ocean is, on average, 4267 m or 14,000 ft deep. These realms and zones are relevant to freshwater lakes as well.
Art Connection
In which of the following regions would you expect to find photosynthetic organisms?
1. the aphotic zone, the neritic zone, the oceanic zone, and the benthic realm
2. the photic zone, the intertidal zone, the neritic zone, and the oceanic zone
3. the photic zone, the abyssal zone, the neritic zone, and the oceanic zone
4. the pelagic realm, the aphotic zone, the neritic zone, and the oceanic zone
Marine Biomes
The ocean is the largest marine biome. It is a continuous body of salt water that is relatively uniform in chemical composition; it is a weak solution of mineral salts and decayed biological matter. Within the ocean, coral reefs are a second kind of marine biome. Estuaries, coastal areas where salt water and fresh water mix, form a third unique marine biome.
Ocean
The physical diversity of the ocean is a significant influence on plants, animals, and other organisms. The ocean is categorized into different zones based on how far light reaches into the water. Each zone has a distinct group of species adapted to the biotic and abiotic conditions particular to that zone.
The intertidal zone, which is the zone between high and low tide, is the oceanic region that is closest to land (Figure \(1\)). Generally, most people think of this portion of the ocean as a sandy beach. In some cases, the intertidal zone is indeed a sandy beach, but it can also be rocky or muddy. The intertidal zone is an extremely variable environment because of tides. Organisms are exposed to air and sunlight at low tide and are underwater most of the time, especially during high tide. Therefore, living things that thrive in the intertidal zone are adapted to being dry for long periods of time. The shore of the intertidal zone is also repeatedly struck by waves, and the organisms found there are adapted to withstand damage from the pounding action of the waves (Figure \(2\)). The exoskeletons of shoreline crustaceans (such as the shore crab, Carcinus maenas) are tough and protect them from desiccation (drying out) and wave damage. Another consequence of the pounding waves is that few algae and plants establish themselves in the constantly moving rocks, sand, or mud.
The neritic zone (Figure \(1\)) extends from the intertidal zone to depths of about 200 m (or 650 ft) at the edge of the continental shelf. Since light can penetrate this depth, photosynthesis can occur in the neritic zone. The water here contains silt and is well-oxygenated, low in pressure, and stable in temperature. Phytoplankton and floating Sargassum (a type of free-floating marine seaweed) provide a habitat for some sea life found in the neritic zone. Zooplankton, protists, small fishes, and shrimp are found in the neritic zone and are the base of the food chain for most of the world’s fisheries.
Beyond the neritic zone is the open ocean area known as the oceanic zone (Figure \(1\)). Within the oceanic zone there is thermal stratification where warm and cold waters mix because of ocean currents. Abundant plankton serve as the base of the food chain for larger animals such as whales and dolphins. Nutrients are scarce and this is a relatively less productive part of the marine biome. When photosynthetic organisms and the protists and animals that feed on them die, their bodies fall to the bottom of the ocean where they remain; unlike freshwater lakes, the open ocean lacks a process for bringing the organic nutrients back up to the surface. The majority of organisms in the aphotic zone include sea cucumbers (phylum Echinodermata) and other organisms that survive on the nutrients contained in the dead bodies of organisms in the photic zone.
Beneath the pelagic zone is the benthic realm, the deepwater region beyond the continental shelf (Figure \(1\)). The bottom of the benthic realm is comprised of sand, silt, and dead organisms. Temperature decreases, remaining above freezing, as water depth increases. This is a nutrient-rich portion of the ocean because of the dead organisms that fall from the upper layers of the ocean. Because of this high level of nutrients, a diversity of fungi, sponges, sea anemones, marine worms, sea stars, fishes, and bacteria exist.
The deepest part of the ocean is the abyssal zone, which is at depths of 4000 m or greater. The abyssal zone (Figure \(1\)) is very cold and has very high pressure, high oxygen content, and low nutrient content. There are a variety of invertebrates and fishes found in this zone, but the abyssal zone does not have plants because of the lack of light. Hydrothermal vents are found primarily in the abyssal zone; chemosynthetic bacteria utilize the hydrogen sulfide and other minerals emitted from the vents. These chemosynthetic bacteria use the hydrogen sulfide as an energy source and serve as the base of the food chain found in the abyssal zone.
Coral Reefs
Coral reefs are ocean ridges formed by marine invertebrates living in warm shallow waters within the photic zone of the ocean. They are found within 30˚ north and south of the equator. The Great Barrier Reef is a well-known reef system located several miles off the northeastern coast of Australia. Other coral reef systems are fringing islands, which are directly adjacent to land, or atolls, which are circular reef systems surrounding a former landmass that is now underwater. The coral organisms (members of phylum Cnidaria) are colonies of saltwater polyps that secrete a calcium carbonate skeleton. These calcium-rich skeletons slowly accumulate, forming the underwater reef (Figure \(3\)). Corals found in shallower waters (at a depth of approximately 60 m or about 200 ft) have a mutualistic relationship with photosynthetic unicellular algae. The relationship provides corals with the majority of the nutrition and the energy they require. The waters in which these corals live are nutritionally poor and, without this mutualism, it would not be possible for large corals to grow. Some corals living in deeper and colder water do not have a mutualistic relationship with algae; these corals attain energy and nutrients using stinging cells on their tentacles to capture prey.
It is estimated that more than 4,000 fish species inhabit coral reefs. These fishes can feed on coral, the cryptofauna (invertebrates found within the calcium carbonate substrate of the coral reefs), or the seaweed and algae that are associated with the coral. In addition, some fish species inhabit the boundaries of a coral reef; these species include predators, herbivores, or planktivores. Predators are animal species that hunt and are carnivores or “flesh eaters.” Herbivores eat plant material, and planktivores eat plankton.
Evolution Connection: Global Decline of Coral Reefs
It takes a long time to build a coral reef. The animals that create coral reefs have evolved over millions of years, continuing to slowly deposit the calcium carbonate that forms their characteristic ocean homes. Bathed in warm tropical waters, the coral animals and their symbiotic algal partners evolved to survive at the upper limit of ocean water temperature.
Together, climate change and human activity pose dual threats to the long-term survival of the world’s coral reefs. As global warming due to fossil fuel emissions raises ocean temperatures, coral reefs are suffering. The excessive warmth causes the reefs to expel their symbiotic, food-producing algae, resulting in a phenomenon known as bleaching. When bleaching occurs, the reefs lose much of their characteristic color as the algae and the coral animals die if loss of the symbiotic zooxanthellae is prolonged.
Rising levels of atmospheric carbon dioxide further threaten the corals in other ways; as CO2 dissolves in ocean waters, it lowers the pH and increases ocean acidity. As acidity increases, it interferes with the calcification that normally occurs as coral animals build their calcium carbonate homes.
When a coral reef begins to die, species diversity plummets as animals lose food and shelter. Coral reefs are also economically important tourist destinations, so the decline of coral reefs poses a serious threat to coastal economies.
Human population growth has damaged corals in other ways, too. As human coastal populations increase, the runoff of sediment and agricultural chemicals has increased, too, causing some of the once-clear tropical waters to become cloudy. At the same time, overfishing of popular fish species has allowed the predator species that eat corals to go unchecked.
Although a rise in global temperatures of 1–2˚C (a conservative scientific projection) in the coming decades may not seem large, it is very significant to this biome. When change occurs rapidly, species can become extinct before evolution leads to new adaptations. Many scientists believe that global warming, with its rapid (in terms of evolutionary time) and inexorable increases in temperature, is tipping the balance beyond the point at which many of the world’s coral reefs can recover.
Estuaries: Where the Ocean Meets Fresh Water
Estuaries are biomes that occur where a source of fresh water, such as a river, meets the ocean. Therefore, both fresh water and salt water are found in the same vicinity; mixing results in a diluted (brackish) saltwater. Estuaries form protected areas where many of the young offspring of crustaceans, mollusks, and fish begin their lives. Salinity is a very important factor that influences the organisms and the adaptations of the organisms found in estuaries. The salinity of estuaries varies and is based on the rate of flow of its freshwater sources. Once or twice a day, high tides bring salt water into the estuary. Low tides occurring at the same frequency reverse the current of salt water.
The short-term and rapid variation in salinity due to the mixing of fresh water and salt water is a difficult physiological challenge for the plants and animals that inhabit estuaries. Many estuarine plant species are halophytes: plants that can tolerate salty conditions. Halophytic plants are adapted to deal with the salinity resulting from saltwater on their roots or from sea spray. In some halophytes, filters in the roots remove the salt from the water that the plant absorbs. Other plants are able to pump oxygen into their roots. Animals, such as mussels and clams (phylum Mollusca), have developed behavioral adaptations that expend a lot of energy to function in this rapidly changing environment. When these animals are exposed to low salinity, they stop feeding, close their shells, and switch from aerobic respiration (in which they use gills) to anaerobic respiration (a process that does not require oxygen). When high tide returns to the estuary, the salinity and oxygen content of the water increases, and these animals open their shells, begin feeding, and return to aerobic respiration.
Freshwater Biomes
Freshwater biomes include lakes and ponds (standing water) as well as rivers and streams (flowing water). They also include wetlands, which will be discussed later. Humans rely on freshwater biomes to provide aquatic resources for drinking water, crop irrigation, sanitation, and industry. These various roles and human benefits are referred to as ecosystem services. Lakes and ponds are found in terrestrial landscapes and are, therefore, connected with abiotic and biotic factors influencing these terrestrial biomes.
Lakes and Ponds
Lakes and ponds can range in area from a few square meters to thousands of square kilometers. Temperature is an important abiotic factor affecting living things found in lakes and ponds. In the summer, thermal stratification of lakes and ponds occurs when the upper layer of water is warmed by the sun and does not mix with deeper, cooler water. Light can penetrate within the photic zone of the lake or pond. Phytoplankton (algae and cyanobacteria) are found here and carry out photosynthesis, providing the base of the food web of lakes and ponds. Zooplankton, such as rotifers and small crustaceans, consume these phytoplankton. At the bottom of lakes and ponds, bacteria in the aphotic zone break down dead organisms that sink to the bottom.
Nitrogen and phosphorus are important limiting nutrients in lakes and ponds. Because of this, they are determining factors in the amount of phytoplankton growth in lakes and ponds. When there is a large input of nitrogen and phosphorus (from sewage and runoff from fertilized lawns and farms, for example), the growth of algae skyrockets, resulting in a large accumulation of algae called an algal bloom. Algal blooms (Figure \(4\)) can become so extensive that they reduce light penetration in water. As a result, the lake or pond becomes aphotic and photosynthetic plants cannot survive. When the algae die and decompose, severe oxygen depletion of the water occurs. Fishes and other organisms that require oxygen are then more likely to die, and resulting dead zones are found across the globe. Lake Erie and the Gulf of Mexico represent freshwater and marine habitats where phosphorus control and storm water runoff pose significant environmental challenges.
Rivers and Streams
Rivers and streams are continuously moving bodies of water that carry large amounts of water from the source, or headwater, to a lake or ocean. The largest rivers include the Nile River in Africa, the Amazon River in South America, and the Mississippi River in North America.
Abiotic features of rivers and streams vary along the length of the river or stream. Streams begin at a point of origin referred to as source water. The source water is usually cold, low in nutrients, and clear. The channel (the width of the river or stream) is narrower than at any other place along the length of the river or stream. Because of this, the current is often faster here than at any other point of the river or stream.
The fast-moving water results in minimal silt accumulation at the bottom of the river or stream; therefore, the water is clear. Photosynthesis here is mostly attributed to algae that are growing on rocks; the swift current inhibits the growth of phytoplankton. An additional input of energy can come from leaves or other organic material that falls into the river or stream from trees and other plants that border the water. When the leaves decompose, the organic material and nutrients in the leaves are returned to the water. Plants and animals have adapted to this fast-moving water. For instance, leeches (phylum Annelida) have elongated bodies and suckers on both ends. These suckers attach to the substrate, keeping the leech anchored in place. Freshwater trout species (phylum Chordata) are an important predator in these fast-moving rivers and streams.
As the river or stream flows away from the source, the width of the channel gradually widens and the current slows. This slow-moving water, caused by the gradient decrease and the volume increase as tributaries unite, has more sedimentation. Phytoplankton can also be suspended in slow-moving water. Therefore, the water will not be as clear as it is near the source. The water is also warmer. Worms (phylum Annelida) and insects (phylum Arthropoda) can be found burrowing into the mud. The higher order predator vertebrates (phylum Chordata) include waterfowl, frogs, and fishes. These predators must find food in these slow moving, sometimes murky, waters and, unlike the trout in the waters at the source, these vertebrates may not be able to use vision as their primary sense to find food. Instead, they are more likely to use taste or chemical cues to find prey.
Wetlands
Wetlands are environments in which the soil is either permanently or periodically saturated with water. Wetlands are different from lakes because wetlands are shallow bodies of water whereas lakes vary in depth. Emergent vegetation consists of wetland plants that are rooted in the soil but have portions of leaves, stems, and flowers extending above the water’s surface. There are several types of wetlands including marshes, swamps, bogs, mudflats, and salt marshes (Figure \(5\)). The three shared characteristics among these types—what makes them wetlands—are their hydrology, hydrophytic vegetation, and hydric soils.
Freshwater marshes and swamps are characterized by slow and steady water flow. Bogs develop in depressions where water flow is low or nonexistent. Bogs usually occur in areas where there is a clay bottom with poor percolation. Percolation is the movement of water through the pores in the soil or rocks. The water found in a bog is stagnant and oxygen depleted because the oxygen that is used during the decomposition of organic matter is not replaced. As the oxygen in the water is depleted, decomposition slows. This leads to organic acids and other acids building up and lowering the pH of the water. At a lower pH, nitrogen becomes unavailable to plants. This creates a challenge for plants because nitrogen is an important limiting resource. Some types of bog plants (such as sundews, pitcher plants, and Venus flytraps) capture insects and extract the nitrogen from their bodies. Bogs have low net primary productivity because the water found in bogs has low levels of nitrogen and oxygen.
Summary
Aquatic ecosystems include both saltwater and freshwater biomes. The abiotic factors important for the structuring of aquatic ecosystems can be different than those seen in terrestrial systems. Sunlight is a driving force behind the structure of forests and also is an important factor in bodies of water, especially those that are very deep, because of the role of photosynthesis in sustaining certain organisms. Density and temperature shape the structure of aquatic systems. Oceans may be thought of as consisting of different zones based on water depth and distance from the shoreline and light penetrance. Different kinds of organisms are adapted to the conditions found in each zone. Coral reefs are unique marine ecosystems that are home to a wide variety of species. Estuaries are found where rivers meet the ocean; their shallow waters provide nourishment and shelter for young crustaceans, mollusks, fishes, and many other species. Freshwater biomes include lakes, ponds, rivers, streams, and wetlands. Bogs are an interesting type of wetland characterized by standing water, lower pH, and a lack of nitrogen.
Art Connections
Figure \(1\): In which of the following regions would you expect to find photosynthetic organisms?
1. the aphotic zone, the neritic zone, the oceanic zone, and the benthic realm
2. the photic zone, the intertidal zone, the neritic zone, and the oceanic zone
3. the photic zone, the abyssal zone, the neritic zone, and the oceanic zone
4. the pelagic realm, the aphotic zone, the neritic zone, and the oceanic zone
Answer
C. Photosynthetic organisms would be found in the photic, abyssal, neritic, and oceanic zones.
Glossary
abyssal zone
deepest part of the ocean at depths of 4000 m or greater
algal bloom
rapid increase of algae in an aquatic system
aphotic zone
part of the ocean where no light penetrates
benthic realm
(also, benthic zone) part of the ocean that extends along the ocean bottom from the shoreline to the deepest parts of the ocean floor
channel
width of a river or stream from one bank to the other bank
coral reef
ocean ridges formed by marine invertebrates living in warm, shallow waters within the photic zone
cryptofauna
invertebrates found within the calcium carbonate substrate of coral reefs
ecosystem services
human benefits and services provided by natural ecosystems
emergent vegetation
wetland plants that are rooted in the soil but have portions of leaves, stems, and flowers extending above the water’s surface
estuary
biomes where a source of fresh water, such as a river, meets the ocean
intertidal zone
part of the ocean that is closest to land; parts extend above the water at low tide
neritic zone
part of the ocean that extends from low tide to the edge of the continental shelf
oceanic zone
part of the ocean that begins offshore where the water measures 200 m deep or deeper
pelagic realm
(also, pelagic zone) open ocean waters that are not close to the bottom or near the shore
photic zone
portion of the ocean that light can penetrate
planktivore
animal species that eats plankton
predator
animal species that hunt and are carnivores or “flesh eaters”
Sargassum
type of free-floating marine seaweed
source water
point of origin of a river or stream | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.03%3A_Freshwater_Habitats/57.3.01%3A_Aquatic_Biomes.txt |
Skills to Develop
• Describe the effects of abiotic factors on the composition of plant and animal communities in aquatic biomes
• Compare and contrast the characteristics of the ocean zones
• Summarize the characteristics of standing water and flowing water freshwater biomes
Abiotic Factors Influencing Aquatic Biomes
Like terrestrial biomes, aquatic biomes are influenced by a series of abiotic factors. The aquatic medium—water— has different physical and chemical properties than air, however. Even if the water in a pond or other body of water is perfectly clear (there are no suspended particles), water, on its own, absorbs light. As one descends into a deep body of water, there will eventually be a depth which the sunlight cannot reach. While there are some abiotic and biotic factors in a terrestrial ecosystem that might obscure light (like fog, dust, or insect swarms), usually these are not permanent features of the environment. The importance of light in aquatic biomes is central to the communities of organisms found in both freshwater and marine ecosystems. In freshwater systems, stratification due to differences in density is perhaps the most critical abiotic factor and is related to the energy aspects of light. The thermal properties of water (rates of heating and cooling) are significant to the function of marine systems and have major impacts on global climate and weather patterns. Marine systems are also influenced by large-scale physical water movements, such as currents; these are less important in most freshwater lakes.
The ocean is categorized by several areas or zones (Figure \(1\)). All of the ocean’s open water is referred to as the pelagic realm (or zone). The benthic realm (or zone) extends along the ocean bottom from the shoreline to the deepest parts of the ocean floor. Within the pelagic realm is the photic zone, which is the portion of the ocean that light can penetrate (approximately 200 m or 650 ft). At depths greater than 200 m, light cannot penetrate; thus, this is referred to as the aphotic zone. The majority of the ocean is aphotic and lacks sufficient light for photosynthesis. The deepest part of the ocean, the Challenger Deep (in the Mariana Trench, located in the western Pacific Ocean), is about 11,000 m (about 6.8 mi) deep. To give some perspective on the depth of this trench, the ocean is, on average, 4267 m or 14,000 ft deep. These realms and zones are relevant to freshwater lakes as well.
Art Connection
In which of the following regions would you expect to find photosynthetic organisms?
1. the aphotic zone, the neritic zone, the oceanic zone, and the benthic realm
2. the photic zone, the intertidal zone, the neritic zone, and the oceanic zone
3. the photic zone, the abyssal zone, the neritic zone, and the oceanic zone
4. the pelagic realm, the aphotic zone, the neritic zone, and the oceanic zone
Marine Biomes
The ocean is the largest marine biome. It is a continuous body of salt water that is relatively uniform in chemical composition; it is a weak solution of mineral salts and decayed biological matter. Within the ocean, coral reefs are a second kind of marine biome. Estuaries, coastal areas where salt water and fresh water mix, form a third unique marine biome.
Ocean
The physical diversity of the ocean is a significant influence on plants, animals, and other organisms. The ocean is categorized into different zones based on how far light reaches into the water. Each zone has a distinct group of species adapted to the biotic and abiotic conditions particular to that zone.
The intertidal zone, which is the zone between high and low tide, is the oceanic region that is closest to land (Figure \(1\)). Generally, most people think of this portion of the ocean as a sandy beach. In some cases, the intertidal zone is indeed a sandy beach, but it can also be rocky or muddy. The intertidal zone is an extremely variable environment because of tides. Organisms are exposed to air and sunlight at low tide and are underwater most of the time, especially during high tide. Therefore, living things that thrive in the intertidal zone are adapted to being dry for long periods of time. The shore of the intertidal zone is also repeatedly struck by waves, and the organisms found there are adapted to withstand damage from the pounding action of the waves (Figure \(2\)). The exoskeletons of shoreline crustaceans (such as the shore crab, Carcinus maenas) are tough and protect them from desiccation (drying out) and wave damage. Another consequence of the pounding waves is that few algae and plants establish themselves in the constantly moving rocks, sand, or mud.
The neritic zone (Figure \(1\)) extends from the intertidal zone to depths of about 200 m (or 650 ft) at the edge of the continental shelf. Since light can penetrate this depth, photosynthesis can occur in the neritic zone. The water here contains silt and is well-oxygenated, low in pressure, and stable in temperature. Phytoplankton and floating Sargassum (a type of free-floating marine seaweed) provide a habitat for some sea life found in the neritic zone. Zooplankton, protists, small fishes, and shrimp are found in the neritic zone and are the base of the food chain for most of the world’s fisheries.
Beyond the neritic zone is the open ocean area known as the oceanic zone (Figure \(1\)). Within the oceanic zone there is thermal stratification where warm and cold waters mix because of ocean currents. Abundant plankton serve as the base of the food chain for larger animals such as whales and dolphins. Nutrients are scarce and this is a relatively less productive part of the marine biome. When photosynthetic organisms and the protists and animals that feed on them die, their bodies fall to the bottom of the ocean where they remain; unlike freshwater lakes, the open ocean lacks a process for bringing the organic nutrients back up to the surface. The majority of organisms in the aphotic zone include sea cucumbers (phylum Echinodermata) and other organisms that survive on the nutrients contained in the dead bodies of organisms in the photic zone.
Beneath the pelagic zone is the benthic realm, the deepwater region beyond the continental shelf (Figure \(1\)). The bottom of the benthic realm is comprised of sand, silt, and dead organisms. Temperature decreases, remaining above freezing, as water depth increases. This is a nutrient-rich portion of the ocean because of the dead organisms that fall from the upper layers of the ocean. Because of this high level of nutrients, a diversity of fungi, sponges, sea anemones, marine worms, sea stars, fishes, and bacteria exist.
The deepest part of the ocean is the abyssal zone, which is at depths of 4000 m or greater. The abyssal zone (Figure \(1\)) is very cold and has very high pressure, high oxygen content, and low nutrient content. There are a variety of invertebrates and fishes found in this zone, but the abyssal zone does not have plants because of the lack of light. Hydrothermal vents are found primarily in the abyssal zone; chemosynthetic bacteria utilize the hydrogen sulfide and other minerals emitted from the vents. These chemosynthetic bacteria use the hydrogen sulfide as an energy source and serve as the base of the food chain found in the abyssal zone.
Coral Reefs
Coral reefs are ocean ridges formed by marine invertebrates living in warm shallow waters within the photic zone of the ocean. They are found within 30˚ north and south of the equator. The Great Barrier Reef is a well-known reef system located several miles off the northeastern coast of Australia. Other coral reef systems are fringing islands, which are directly adjacent to land, or atolls, which are circular reef systems surrounding a former landmass that is now underwater. The coral organisms (members of phylum Cnidaria) are colonies of saltwater polyps that secrete a calcium carbonate skeleton. These calcium-rich skeletons slowly accumulate, forming the underwater reef (Figure \(3\)). Corals found in shallower waters (at a depth of approximately 60 m or about 200 ft) have a mutualistic relationship with photosynthetic unicellular algae. The relationship provides corals with the majority of the nutrition and the energy they require. The waters in which these corals live are nutritionally poor and, without this mutualism, it would not be possible for large corals to grow. Some corals living in deeper and colder water do not have a mutualistic relationship with algae; these corals attain energy and nutrients using stinging cells on their tentacles to capture prey.
It is estimated that more than 4,000 fish species inhabit coral reefs. These fishes can feed on coral, the cryptofauna (invertebrates found within the calcium carbonate substrate of the coral reefs), or the seaweed and algae that are associated with the coral. In addition, some fish species inhabit the boundaries of a coral reef; these species include predators, herbivores, or planktivores. Predators are animal species that hunt and are carnivores or “flesh eaters.” Herbivores eat plant material, and planktivores eat plankton.
Evolution Connection: Global Decline of Coral Reefs
It takes a long time to build a coral reef. The animals that create coral reefs have evolved over millions of years, continuing to slowly deposit the calcium carbonate that forms their characteristic ocean homes. Bathed in warm tropical waters, the coral animals and their symbiotic algal partners evolved to survive at the upper limit of ocean water temperature.
Together, climate change and human activity pose dual threats to the long-term survival of the world’s coral reefs. As global warming due to fossil fuel emissions raises ocean temperatures, coral reefs are suffering. The excessive warmth causes the reefs to expel their symbiotic, food-producing algae, resulting in a phenomenon known as bleaching. When bleaching occurs, the reefs lose much of their characteristic color as the algae and the coral animals die if loss of the symbiotic zooxanthellae is prolonged.
Rising levels of atmospheric carbon dioxide further threaten the corals in other ways; as CO2 dissolves in ocean waters, it lowers the pH and increases ocean acidity. As acidity increases, it interferes with the calcification that normally occurs as coral animals build their calcium carbonate homes.
When a coral reef begins to die, species diversity plummets as animals lose food and shelter. Coral reefs are also economically important tourist destinations, so the decline of coral reefs poses a serious threat to coastal economies.
Human population growth has damaged corals in other ways, too. As human coastal populations increase, the runoff of sediment and agricultural chemicals has increased, too, causing some of the once-clear tropical waters to become cloudy. At the same time, overfishing of popular fish species has allowed the predator species that eat corals to go unchecked.
Although a rise in global temperatures of 1–2˚C (a conservative scientific projection) in the coming decades may not seem large, it is very significant to this biome. When change occurs rapidly, species can become extinct before evolution leads to new adaptations. Many scientists believe that global warming, with its rapid (in terms of evolutionary time) and inexorable increases in temperature, is tipping the balance beyond the point at which many of the world’s coral reefs can recover.
Estuaries: Where the Ocean Meets Fresh Water
Estuaries are biomes that occur where a source of fresh water, such as a river, meets the ocean. Therefore, both fresh water and salt water are found in the same vicinity; mixing results in a diluted (brackish) saltwater. Estuaries form protected areas where many of the young offspring of crustaceans, mollusks, and fish begin their lives. Salinity is a very important factor that influences the organisms and the adaptations of the organisms found in estuaries. The salinity of estuaries varies and is based on the rate of flow of its freshwater sources. Once or twice a day, high tides bring salt water into the estuary. Low tides occurring at the same frequency reverse the current of salt water.
The short-term and rapid variation in salinity due to the mixing of fresh water and salt water is a difficult physiological challenge for the plants and animals that inhabit estuaries. Many estuarine plant species are halophytes: plants that can tolerate salty conditions. Halophytic plants are adapted to deal with the salinity resulting from saltwater on their roots or from sea spray. In some halophytes, filters in the roots remove the salt from the water that the plant absorbs. Other plants are able to pump oxygen into their roots. Animals, such as mussels and clams (phylum Mollusca), have developed behavioral adaptations that expend a lot of energy to function in this rapidly changing environment. When these animals are exposed to low salinity, they stop feeding, close their shells, and switch from aerobic respiration (in which they use gills) to anaerobic respiration (a process that does not require oxygen). When high tide returns to the estuary, the salinity and oxygen content of the water increases, and these animals open their shells, begin feeding, and return to aerobic respiration.
Freshwater Biomes
Freshwater biomes include lakes and ponds (standing water) as well as rivers and streams (flowing water). They also include wetlands, which will be discussed later. Humans rely on freshwater biomes to provide aquatic resources for drinking water, crop irrigation, sanitation, and industry. These various roles and human benefits are referred to as ecosystem services. Lakes and ponds are found in terrestrial landscapes and are, therefore, connected with abiotic and biotic factors influencing these terrestrial biomes.
Lakes and Ponds
Lakes and ponds can range in area from a few square meters to thousands of square kilometers. Temperature is an important abiotic factor affecting living things found in lakes and ponds. In the summer, thermal stratification of lakes and ponds occurs when the upper layer of water is warmed by the sun and does not mix with deeper, cooler water. Light can penetrate within the photic zone of the lake or pond. Phytoplankton (algae and cyanobacteria) are found here and carry out photosynthesis, providing the base of the food web of lakes and ponds. Zooplankton, such as rotifers and small crustaceans, consume these phytoplankton. At the bottom of lakes and ponds, bacteria in the aphotic zone break down dead organisms that sink to the bottom.
Nitrogen and phosphorus are important limiting nutrients in lakes and ponds. Because of this, they are determining factors in the amount of phytoplankton growth in lakes and ponds. When there is a large input of nitrogen and phosphorus (from sewage and runoff from fertilized lawns and farms, for example), the growth of algae skyrockets, resulting in a large accumulation of algae called an algal bloom. Algal blooms (Figure \(4\)) can become so extensive that they reduce light penetration in water. As a result, the lake or pond becomes aphotic and photosynthetic plants cannot survive. When the algae die and decompose, severe oxygen depletion of the water occurs. Fishes and other organisms that require oxygen are then more likely to die, and resulting dead zones are found across the globe. Lake Erie and the Gulf of Mexico represent freshwater and marine habitats where phosphorus control and storm water runoff pose significant environmental challenges.
Rivers and Streams
Rivers and streams are continuously moving bodies of water that carry large amounts of water from the source, or headwater, to a lake or ocean. The largest rivers include the Nile River in Africa, the Amazon River in South America, and the Mississippi River in North America.
Abiotic features of rivers and streams vary along the length of the river or stream. Streams begin at a point of origin referred to as source water. The source water is usually cold, low in nutrients, and clear. The channel (the width of the river or stream) is narrower than at any other place along the length of the river or stream. Because of this, the current is often faster here than at any other point of the river or stream.
The fast-moving water results in minimal silt accumulation at the bottom of the river or stream; therefore, the water is clear. Photosynthesis here is mostly attributed to algae that are growing on rocks; the swift current inhibits the growth of phytoplankton. An additional input of energy can come from leaves or other organic material that falls into the river or stream from trees and other plants that border the water. When the leaves decompose, the organic material and nutrients in the leaves are returned to the water. Plants and animals have adapted to this fast-moving water. For instance, leeches (phylum Annelida) have elongated bodies and suckers on both ends. These suckers attach to the substrate, keeping the leech anchored in place. Freshwater trout species (phylum Chordata) are an important predator in these fast-moving rivers and streams.
As the river or stream flows away from the source, the width of the channel gradually widens and the current slows. This slow-moving water, caused by the gradient decrease and the volume increase as tributaries unite, has more sedimentation. Phytoplankton can also be suspended in slow-moving water. Therefore, the water will not be as clear as it is near the source. The water is also warmer. Worms (phylum Annelida) and insects (phylum Arthropoda) can be found burrowing into the mud. The higher order predator vertebrates (phylum Chordata) include waterfowl, frogs, and fishes. These predators must find food in these slow moving, sometimes murky, waters and, unlike the trout in the waters at the source, these vertebrates may not be able to use vision as their primary sense to find food. Instead, they are more likely to use taste or chemical cues to find prey.
Wetlands
Wetlands are environments in which the soil is either permanently or periodically saturated with water. Wetlands are different from lakes because wetlands are shallow bodies of water whereas lakes vary in depth. Emergent vegetation consists of wetland plants that are rooted in the soil but have portions of leaves, stems, and flowers extending above the water’s surface. There are several types of wetlands including marshes, swamps, bogs, mudflats, and salt marshes (Figure \(5\)). The three shared characteristics among these types—what makes them wetlands—are their hydrology, hydrophytic vegetation, and hydric soils.
Freshwater marshes and swamps are characterized by slow and steady water flow. Bogs develop in depressions where water flow is low or nonexistent. Bogs usually occur in areas where there is a clay bottom with poor percolation. Percolation is the movement of water through the pores in the soil or rocks. The water found in a bog is stagnant and oxygen depleted because the oxygen that is used during the decomposition of organic matter is not replaced. As the oxygen in the water is depleted, decomposition slows. This leads to organic acids and other acids building up and lowering the pH of the water. At a lower pH, nitrogen becomes unavailable to plants. This creates a challenge for plants because nitrogen is an important limiting resource. Some types of bog plants (such as sundews, pitcher plants, and Venus flytraps) capture insects and extract the nitrogen from their bodies. Bogs have low net primary productivity because the water found in bogs has low levels of nitrogen and oxygen.
Summary
Aquatic ecosystems include both saltwater and freshwater biomes. The abiotic factors important for the structuring of aquatic ecosystems can be different than those seen in terrestrial systems. Sunlight is a driving force behind the structure of forests and also is an important factor in bodies of water, especially those that are very deep, because of the role of photosynthesis in sustaining certain organisms. Density and temperature shape the structure of aquatic systems. Oceans may be thought of as consisting of different zones based on water depth and distance from the shoreline and light penetrance. Different kinds of organisms are adapted to the conditions found in each zone. Coral reefs are unique marine ecosystems that are home to a wide variety of species. Estuaries are found where rivers meet the ocean; their shallow waters provide nourishment and shelter for young crustaceans, mollusks, fishes, and many other species. Freshwater biomes include lakes, ponds, rivers, streams, and wetlands. Bogs are an interesting type of wetland characterized by standing water, lower pH, and a lack of nitrogen.
Art Connections
Figure \(1\): In which of the following regions would you expect to find photosynthetic organisms?
1. the aphotic zone, the neritic zone, the oceanic zone, and the benthic realm
2. the photic zone, the intertidal zone, the neritic zone, and the oceanic zone
3. the photic zone, the abyssal zone, the neritic zone, and the oceanic zone
4. the pelagic realm, the aphotic zone, the neritic zone, and the oceanic zone
Answer
C. Photosynthetic organisms would be found in the photic, abyssal, neritic, and oceanic zones.
Glossary
abyssal zone
deepest part of the ocean at depths of 4000 m or greater
algal bloom
rapid increase of algae in an aquatic system
aphotic zone
part of the ocean where no light penetrates
benthic realm
(also, benthic zone) part of the ocean that extends along the ocean bottom from the shoreline to the deepest parts of the ocean floor
channel
width of a river or stream from one bank to the other bank
coral reef
ocean ridges formed by marine invertebrates living in warm, shallow waters within the photic zone
cryptofauna
invertebrates found within the calcium carbonate substrate of coral reefs
ecosystem services
human benefits and services provided by natural ecosystems
emergent vegetation
wetland plants that are rooted in the soil but have portions of leaves, stems, and flowers extending above the water’s surface
estuary
biomes where a source of fresh water, such as a river, meets the ocean
intertidal zone
part of the ocean that is closest to land; parts extend above the water at low tide
neritic zone
part of the ocean that extends from low tide to the edge of the continental shelf
oceanic zone
part of the ocean that begins offshore where the water measures 200 m deep or deeper
pelagic realm
(also, pelagic zone) open ocean waters that are not close to the bottom or near the shore
photic zone
portion of the ocean that light can penetrate
planktivore
animal species that eats plankton
predator
animal species that hunt and are carnivores or “flesh eaters”
Sargassum
type of free-floating marine seaweed
source water
point of origin of a river or stream | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.04%3A_Marine_Habitats.txt |
Skills to Develop
• Define global climate change
• Summarize the effects of the Industrial Revolution on global atmospheric carbon dioxide concentration
• Describe three natural factors affecting long-term global climate
• List two or more greenhouse gases and describe their role in the greenhouse effect
All biomes are universally affected by global conditions, such as climate, that ultimately shape each biome’s environment. Scientists who study climate have noted a series of marked changes that have gradually become increasingly evident during the last sixty years. Global climate change is the term used to describe altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide.
Climate and Weather
A common misconception about global climate change is that a specific weather event occurring in a particular region (for example, a very cool week in June in central Indiana) is evidence of global climate change. However, a cold week in June is a weather-related event and not a climate-related one. These misconceptions often arise because of confusion over the terms climate and weather.
Climate refers to the long-term, predictable atmospheric conditions of a specific area. The climate of a biome is characterized by having consistent temperature and annual rainfall ranges. Climate does not address the amount of rain that fell on one particular day in a biome or the colder-than-average temperatures that occurred on one day. In contrast, weather refers to the conditions of the atmosphere during a short period of time. Weather forecasts are usually made for 48-hour cycles. Long-range weather forecasts are available but can be unreliable.
To better understand the difference between climate and weather, imagine that you are planning an outdoor event in northern Wisconsin. You would be thinking about climate when you plan the event in the summer rather than the winter because you have long-term knowledge that any given Saturday in the months of May to August would be a better choice for an outdoor event in Wisconsin than any given Saturday in January. However, you cannot determine the specific day that the event should be held on because it is difficult to accurately predict the weather on a specific day. Climate can be considered “average” weather.
Global Climate Change
Climate change can be understood by approaching three areas of study:
• current and past global climate change
• causes of past and present-day global climate change
• ancient and current results of climate change
It is helpful to keep these three different aspects of climate change clearly separated when consuming media reports about global climate change. It is common for reports and discussions about global climate change to confuse the data showing that Earth’s climate is changing with the factors that drive this climate change.
Evidence for Global Climate Change
Since scientists cannot go back in time to directly measure climatic variables, such as average temperature and precipitation, they must instead indirectly measure temperature. To do this, scientists rely on historical evidence of Earth’s past climate.
Antarctic ice cores are a key example of such evidence. These ice cores are samples of polar ice obtained by means of drills that reach thousands of meters into ice sheets or high mountain glaciers. Viewing the ice cores is like traveling backwards through time; the deeper the sample, the earlier the time period. Trapped within the ice are bubbles of air and other biological evidence that can reveal temperature and carbon dioxide data. Antarctic ice cores have been collected and analyzed to indirectly estimate the temperature of the Earth over the past 400,000 years (Figure \(1\)a). The 0 °C on this graph refers to the long-term average. Temperatures that are greater than 0 °C exceed Earth’s long-term average temperature. Conversely, temperatures that are less than 0 °C are less than Earth’s average temperature. This figure shows that there have been periodic cycles of increasing and decreasing temperature.
Before the late 1800s, the Earth has been as much as 9 °C cooler and about 3 °C warmer. Note that the graph in Figure \(1\)b shows that the atmospheric concentration of carbon dioxide has also risen and fallen in periodic cycles; note the relationship between carbon dioxide concentration and temperature. Figure \(1\)b shows that carbon dioxide levels in the atmosphere have historically cycled between 180 and 300 parts per million (ppm) by volume.
Figure \(1\)a does not show the last 2,000 years with enough detail to compare the changes of Earth’s temperature during the last 400,000 years with the temperature change that has occurred in the more recent past. Two significant temperature anomalies, or irregularities, have occurred in the last 2000 years. These are the Medieval Climate Anomaly (or the Medieval Warm Period) and the Little Ice Age. A third temperature anomaly aligns with the Industrial Era. The Medieval Climate Anomaly occurred between 900 and 1300 AD. During this time period, many climate scientists think that slightly warmer weather conditions prevailed in many parts of the world; the higher-than-average temperature changes varied between 0.10 °C and 0.20 °C above the norm. Although 0.10 °C does not seem large enough to produce any noticeable change, it did free seas of ice. Because of this warming, the Vikings were able to colonize Greenland.
The Little Ice Age was a cold period that occurred between 1550 AD and 1850 AD. During this time, a slight cooling of a little less than 1 °C was observed in North America, Europe, and possibly other areas of the Earth. This 1 °C change in global temperature is a seemingly small deviation in temperature (as was observed during the Medieval Climate Anomaly); however, it also resulted in noticeable changes. Historical accounts reveal a time of exceptionally harsh winters with much snow and frost.
The Industrial Revolution, which began around 1750, was characterized by changes in much of human society. Advances in agriculture increased the food supply, which improved the standard of living for people in Europe and the United States. New technologies were invented and provided jobs and cheaper goods. These new technologies were powered using fossil fuels, especially coal. The Industrial Revolution starting in the early nineteenth century ushered in the beginning of the Industrial Era. When a fossil fuel is burned, carbon dioxide is released. With the beginning of the Industrial Era, atmospheric carbon dioxide began to rise (Figure \(2\)).
Current and Past Drivers of Global Climate Change
Since it is not possible to go back in time to directly observe and measure climate, scientists use indirect evidence to determine the drivers, or factors, that may be responsible for climate change. The indirect evidence includes data collected using ice cores, boreholes (a narrow shaft bored into the ground), tree rings, glacier lengths, pollen remains, and ocean sediments. The data shows a correlation between the timing of temperature changes and drivers of climate change: before the Industrial Era (pre-1780), there were three drivers of climate change that were not related to human activity or atmospheric gases. The first of these is the Milankovitch cycles. The Milankovitch cycles describe the effects of slight changes in the Earth’s orbit on Earth’s climate. The length of the Milankovitch cycles ranges between 19,000 and 100,000 years. In other words, one could expect to see some predictable changes in the Earth’s climate associated with changes in the Earth’s orbit at a minimum of every 19,000 years.
The variation in the sun’s intensity is the second natural factor responsible for climate change. Solar intensity is the amount of solar power or energy the sun emits in a given amount of time. There is a direct relationship between solar intensity and temperature. As solar intensity increases (or decreases), the Earth’s temperature correspondingly increases (or decreases). Changes in solar intensity have been proposed as one of several possible explanations for the Little Ice Age.
Finally, volcanic eruptions are a third natural driver of climate change. Volcanic eruptions can last a few days, but the solids and gases released during an eruption can influence the climate over a period of a few years, causing short-term climate changes. The gases and solids released by volcanic eruptions can include carbon dioxide, water vapor, sulfur dioxide, hydrogen sulfide, hydrogen, and carbon monoxide. Generally, volcanic eruptions cool the climate. This occurred in 1783 when volcanos in Iceland erupted and caused the release of large volumes of sulfuric oxide. This led to haze-effect cooling, a global phenomenon that occurs when dust, ash, or other suspended particles block out sunlight and trigger lower global temperatures as a result; haze-effect cooling usually extends for one or more years. In Europe and North America, haze-effect cooling produced some of the lowest average winter temperatures on record in 1783 and 1784.
Greenhouse gases are probably the most significant drivers of the climate. When heat energy from the sun strikes the Earth, gases known as greenhouse gases trap the heat in the atmosphere, as do the glass panes of a greenhouse keep heat from escaping. The greenhouse gases that affect Earth include carbon dioxide, methane, water vapor, nitrous oxide, and ozone. Approximately half of the radiation from the sun passes through these gases in the atmosphere and strikes the Earth. This radiation is converted into thermal radiation on the Earth’s surface, and then a portion of that energy is re-radiated back into the atmosphere. Greenhouse gases, however, reflect much of the thermal energy back to the Earth’s surface. The more greenhouse gases there are in the atmosphere, the more thermal energy is reflected back to the Earth’s surface. Greenhouse gases absorb and emit radiation and are an important factor in the greenhouse effect: the warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere.
Evidence supports the relationship between atmospheric concentrations of carbon dioxide and temperature: as carbon dioxide rises, global temperature rises. Since 1950, the concentration of atmospheric carbon dioxide has increased from about 280 ppm to 382 ppm in 2006. In 2011, the atmospheric carbon dioxide concentration was 392 ppm. However, the planet would not be inhabitable by current life forms if water vapor did not produce its drastic greenhouse warming effect.
Scientists look at patterns in data and try to explain differences or deviations from these patterns. The atmospheric carbon dioxide data reveal a historical pattern of carbon dioxide increasing and decreasing, cycling between a low of 180 ppm and a high of 300 ppm. Scientists have concluded that it took around 50,000 years for the atmospheric carbon dioxide level to increase from its low minimum concentration to its higher maximum concentration. However, starting recently, atmospheric carbon dioxide concentrations have increased beyond the historical maximum of 300 ppm. The current increases in atmospheric carbon dioxide have happened very quickly—in a matter of hundreds of years rather than thousands of years. What is the reason for this difference in the rate of change and the amount of increase in carbon dioxide? A key factor that must be recognized when comparing the historical data and the current data is the presence of modern human society; no other driver of climate change has yielded changes in atmospheric carbon dioxide levels at this rate or to this magnitude.
Human activity releases carbon dioxide and methane, two of the most important greenhouse gases, into the atmosphere in several ways. The primary mechanism that releases carbon dioxide is the burning of fossil fuels, such as gasoline, coal, and natural gas (Figure \(3\)). Deforestation, cement manufacture, animal agriculture, the clearing of land, and the burning of forests are other human activities that release carbon dioxide. Methane (CH4) is produced when bacteria break down organic matter under anaerobic conditions. Anaerobic conditions can happen when organic matter is trapped underwater (such as in rice paddies) or in the intestines of herbivores. Methane can also be released from natural gas fields and the decomposition that occurs in landfills. Another source of methane is the melting of clathrates. Clathrates are frozen chunks of ice and methane found at the bottom of the ocean. When water warms, these chunks of ice melt and methane is released. As the ocean’s water temperature increases, the rate at which clathrates melt is increasing, releasing even more methane. This leads to increased levels of methane in the atmosphere, which further accelerates the rate of global warming. This is an example of the positive feedback loop that is leading to the rapid rate of increase of global temperatures.
Documented Results of Climate Change: Past and Present
Scientists have geological evidence of the consequences of long-ago climate change. Modern-day phenomena such as retreating glaciers and melting polar ice cause a continual rise in sea level. Meanwhile, changes in climate can negatively affect organisms.
Geological Climate Change
Global warming has been associated with at least one planet-wide extinction event during the geological past. The Permian extinction event occurred about 251 million years ago toward the end of the roughly 50-million-year-long geological time span known as the Permian period. This geologic time period was one of the three warmest periods in Earth’s geologic history. Scientists estimate that approximately 70 percent of the terrestrial plant and animal species and 84 percent of marine species became extinct, vanishing forever near the end of the Permian period. Organisms that had adapted to wet and warm climatic conditions, such as annual rainfall of 300–400 cm (118–157 in) and 20 °C–30 °C (68 °F–86 °F) in the tropical wet forest, may not have been able to survive the Permian climate change.
Link to Learning
Watch this NASA video to discover the mixed effects of global warming on plant growth. While scientists found that warmer temperatures in the 1980s and 1990s caused an increase in plant productivity, this advantage has since been counteracted by more frequent droughts.
Present Climate Change
A number of global events have occurred that may be attributed to climate change during our lifetimes. Glacier National Park in Montana is undergoing the retreat of many of its glaciers, a phenomenon known as glacier recession. In 1850, the area contained approximately 150 glaciers. By 2010, however, the park contained only about 24 glaciers greater than 25 acres in size. One of these glaciers is the Grinnell Glacier (Figure \(4\)) at Mount Gould. Between 1966 and 2005, the size of Grinnell Glacier shrank by 40 percent. Similarly, the mass of the ice sheets in Greenland and the Antarctic is decreasing: Greenland lost 150–250 km3 of ice per year between 2002 and 2006. In addition, the size and thickness of the Arctic sea ice is decreasing.
This loss of ice is leading to increases in the global sea level. On average, the sea is rising at a rate of 1.8 mm per year. However, between 1993 and 2010 the rate of sea level increase ranged between 2.9 and 3.4 mm per year. A variety of factors affect the volume of water in the ocean, including the temperature of the water (the density of water is related to its temperature) and the amount of water found in rivers, lakes, glaciers, polar ice caps, and sea ice. As glaciers and polar ice caps melt, there is a significant contribution of liquid water that was previously frozen.
In addition to some abiotic conditions changing in response to climate change, many organisms are also being affected by the changes in temperature. Temperature and precipitation play key roles in determining the geographic distribution and phenology of plants and animals. (Phenology is the study of the effects of climatic conditions on the timing of periodic lifecycle events, such as flowering in plants or migration in birds.) Researchers have shown that 385 plant species in Great Britain are flowering 4.5 days sooner than was recorded earlier during the previous 40 years. In addition, insect-pollinated species were more likely to flower earlier than wind-pollinated species. The impact of changes in flowering date would be mitigated if the insect pollinators emerged earlier. This mismatched timing of plants and pollinators could result in injurious ecosystem effects because, for continued survival, insect-pollinated plants must flower when their pollinators are present.
Summary
The Earth has gone through periodic cycles of increases and decreases in temperature. During the past 2000 years, the Medieval Climate Anomaly was a warmer period, while the Little Ice Age was unusually cool. Both of these irregularities can be explained by natural causes of changes in climate, and, although the temperature changes were small, they had significant effects. Natural drivers of climate change include Milankovitch cycles, changes in solar activity, and volcanic eruptions. None of these factors, however, leads to rapid increases in global temperature or sustained increases in carbon dioxide. The burning of fossil fuels is an important source of greenhouse gases, which plays a major role in the greenhouse effect. Long ago, global warming resulted in the Permian extinction: a large-scale extinction event that is documented in the fossil record. Currently, modern-day climate change is associated with the increased melting of glaciers and polar ice sheets, resulting in a gradual increase in sea level. Plants and animals can also be affected by global climate change when the timing of seasonal events, such as flowering or pollination, is affected by global warming.
Glossary
clathrates
frozen chunks of ice and methane found at the bottom of the ocean
climate
long-term, predictable atmospheric conditions present in a specific area
global climate change
altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide
greenhouse effect
warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere
greenhouse gases
atmospheric gases such as carbon dioxide and methane that absorb and emit radiation, thus trapping heat in Earth’s atmosphere
haze-effect cooling
effect of the gases and solids from a volcanic eruption on global climate
Milankovitch cycles
cyclic changes in the Earth's orbit that may affect climate
solar intensity
amount of solar power energy the sun emits in a given amount of time
weather
conditions of the atmosphere during a short period of time
57.05: Human Impacts of the Biosphere- Pollution and Resource Depletion
Skills to Develop
• Define global climate change
• Summarize the effects of the Industrial Revolution on global atmospheric carbon dioxide concentration
• Describe three natural factors affecting long-term global climate
• List two or more greenhouse gases and describe their role in the greenhouse effect
All biomes are universally affected by global conditions, such as climate, that ultimately shape each biome’s environment. Scientists who study climate have noted a series of marked changes that have gradually become increasingly evident during the last sixty years. Global climate change is the term used to describe altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide.
Climate and Weather
A common misconception about global climate change is that a specific weather event occurring in a particular region (for example, a very cool week in June in central Indiana) is evidence of global climate change. However, a cold week in June is a weather-related event and not a climate-related one. These misconceptions often arise because of confusion over the terms climate and weather.
Climate refers to the long-term, predictable atmospheric conditions of a specific area. The climate of a biome is characterized by having consistent temperature and annual rainfall ranges. Climate does not address the amount of rain that fell on one particular day in a biome or the colder-than-average temperatures that occurred on one day. In contrast, weather refers to the conditions of the atmosphere during a short period of time. Weather forecasts are usually made for 48-hour cycles. Long-range weather forecasts are available but can be unreliable.
To better understand the difference between climate and weather, imagine that you are planning an outdoor event in northern Wisconsin. You would be thinking about climate when you plan the event in the summer rather than the winter because you have long-term knowledge that any given Saturday in the months of May to August would be a better choice for an outdoor event in Wisconsin than any given Saturday in January. However, you cannot determine the specific day that the event should be held on because it is difficult to accurately predict the weather on a specific day. Climate can be considered “average” weather.
Global Climate Change
Climate change can be understood by approaching three areas of study:
• current and past global climate change
• causes of past and present-day global climate change
• ancient and current results of climate change
It is helpful to keep these three different aspects of climate change clearly separated when consuming media reports about global climate change. It is common for reports and discussions about global climate change to confuse the data showing that Earth’s climate is changing with the factors that drive this climate change.
Evidence for Global Climate Change
Since scientists cannot go back in time to directly measure climatic variables, such as average temperature and precipitation, they must instead indirectly measure temperature. To do this, scientists rely on historical evidence of Earth’s past climate.
Antarctic ice cores are a key example of such evidence. These ice cores are samples of polar ice obtained by means of drills that reach thousands of meters into ice sheets or high mountain glaciers. Viewing the ice cores is like traveling backwards through time; the deeper the sample, the earlier the time period. Trapped within the ice are bubbles of air and other biological evidence that can reveal temperature and carbon dioxide data. Antarctic ice cores have been collected and analyzed to indirectly estimate the temperature of the Earth over the past 400,000 years (Figure \(1\)a). The 0 °C on this graph refers to the long-term average. Temperatures that are greater than 0 °C exceed Earth’s long-term average temperature. Conversely, temperatures that are less than 0 °C are less than Earth’s average temperature. This figure shows that there have been periodic cycles of increasing and decreasing temperature.
Before the late 1800s, the Earth has been as much as 9 °C cooler and about 3 °C warmer. Note that the graph in Figure \(1\)b shows that the atmospheric concentration of carbon dioxide has also risen and fallen in periodic cycles; note the relationship between carbon dioxide concentration and temperature. Figure \(1\)b shows that carbon dioxide levels in the atmosphere have historically cycled between 180 and 300 parts per million (ppm) by volume.
Figure \(1\)a does not show the last 2,000 years with enough detail to compare the changes of Earth’s temperature during the last 400,000 years with the temperature change that has occurred in the more recent past. Two significant temperature anomalies, or irregularities, have occurred in the last 2000 years. These are the Medieval Climate Anomaly (or the Medieval Warm Period) and the Little Ice Age. A third temperature anomaly aligns with the Industrial Era. The Medieval Climate Anomaly occurred between 900 and 1300 AD. During this time period, many climate scientists think that slightly warmer weather conditions prevailed in many parts of the world; the higher-than-average temperature changes varied between 0.10 °C and 0.20 °C above the norm. Although 0.10 °C does not seem large enough to produce any noticeable change, it did free seas of ice. Because of this warming, the Vikings were able to colonize Greenland.
The Little Ice Age was a cold period that occurred between 1550 AD and 1850 AD. During this time, a slight cooling of a little less than 1 °C was observed in North America, Europe, and possibly other areas of the Earth. This 1 °C change in global temperature is a seemingly small deviation in temperature (as was observed during the Medieval Climate Anomaly); however, it also resulted in noticeable changes. Historical accounts reveal a time of exceptionally harsh winters with much snow and frost.
The Industrial Revolution, which began around 1750, was characterized by changes in much of human society. Advances in agriculture increased the food supply, which improved the standard of living for people in Europe and the United States. New technologies were invented and provided jobs and cheaper goods. These new technologies were powered using fossil fuels, especially coal. The Industrial Revolution starting in the early nineteenth century ushered in the beginning of the Industrial Era. When a fossil fuel is burned, carbon dioxide is released. With the beginning of the Industrial Era, atmospheric carbon dioxide began to rise (Figure \(2\)).
Current and Past Drivers of Global Climate Change
Since it is not possible to go back in time to directly observe and measure climate, scientists use indirect evidence to determine the drivers, or factors, that may be responsible for climate change. The indirect evidence includes data collected using ice cores, boreholes (a narrow shaft bored into the ground), tree rings, glacier lengths, pollen remains, and ocean sediments. The data shows a correlation between the timing of temperature changes and drivers of climate change: before the Industrial Era (pre-1780), there were three drivers of climate change that were not related to human activity or atmospheric gases. The first of these is the Milankovitch cycles. The Milankovitch cycles describe the effects of slight changes in the Earth’s orbit on Earth’s climate. The length of the Milankovitch cycles ranges between 19,000 and 100,000 years. In other words, one could expect to see some predictable changes in the Earth’s climate associated with changes in the Earth’s orbit at a minimum of every 19,000 years.
The variation in the sun’s intensity is the second natural factor responsible for climate change. Solar intensity is the amount of solar power or energy the sun emits in a given amount of time. There is a direct relationship between solar intensity and temperature. As solar intensity increases (or decreases), the Earth’s temperature correspondingly increases (or decreases). Changes in solar intensity have been proposed as one of several possible explanations for the Little Ice Age.
Finally, volcanic eruptions are a third natural driver of climate change. Volcanic eruptions can last a few days, but the solids and gases released during an eruption can influence the climate over a period of a few years, causing short-term climate changes. The gases and solids released by volcanic eruptions can include carbon dioxide, water vapor, sulfur dioxide, hydrogen sulfide, hydrogen, and carbon monoxide. Generally, volcanic eruptions cool the climate. This occurred in 1783 when volcanos in Iceland erupted and caused the release of large volumes of sulfuric oxide. This led to haze-effect cooling, a global phenomenon that occurs when dust, ash, or other suspended particles block out sunlight and trigger lower global temperatures as a result; haze-effect cooling usually extends for one or more years. In Europe and North America, haze-effect cooling produced some of the lowest average winter temperatures on record in 1783 and 1784.
Greenhouse gases are probably the most significant drivers of the climate. When heat energy from the sun strikes the Earth, gases known as greenhouse gases trap the heat in the atmosphere, as do the glass panes of a greenhouse keep heat from escaping. The greenhouse gases that affect Earth include carbon dioxide, methane, water vapor, nitrous oxide, and ozone. Approximately half of the radiation from the sun passes through these gases in the atmosphere and strikes the Earth. This radiation is converted into thermal radiation on the Earth’s surface, and then a portion of that energy is re-radiated back into the atmosphere. Greenhouse gases, however, reflect much of the thermal energy back to the Earth’s surface. The more greenhouse gases there are in the atmosphere, the more thermal energy is reflected back to the Earth’s surface. Greenhouse gases absorb and emit radiation and are an important factor in the greenhouse effect: the warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere.
Evidence supports the relationship between atmospheric concentrations of carbon dioxide and temperature: as carbon dioxide rises, global temperature rises. Since 1950, the concentration of atmospheric carbon dioxide has increased from about 280 ppm to 382 ppm in 2006. In 2011, the atmospheric carbon dioxide concentration was 392 ppm. However, the planet would not be inhabitable by current life forms if water vapor did not produce its drastic greenhouse warming effect.
Scientists look at patterns in data and try to explain differences or deviations from these patterns. The atmospheric carbon dioxide data reveal a historical pattern of carbon dioxide increasing and decreasing, cycling between a low of 180 ppm and a high of 300 ppm. Scientists have concluded that it took around 50,000 years for the atmospheric carbon dioxide level to increase from its low minimum concentration to its higher maximum concentration. However, starting recently, atmospheric carbon dioxide concentrations have increased beyond the historical maximum of 300 ppm. The current increases in atmospheric carbon dioxide have happened very quickly—in a matter of hundreds of years rather than thousands of years. What is the reason for this difference in the rate of change and the amount of increase in carbon dioxide? A key factor that must be recognized when comparing the historical data and the current data is the presence of modern human society; no other driver of climate change has yielded changes in atmospheric carbon dioxide levels at this rate or to this magnitude.
Human activity releases carbon dioxide and methane, two of the most important greenhouse gases, into the atmosphere in several ways. The primary mechanism that releases carbon dioxide is the burning of fossil fuels, such as gasoline, coal, and natural gas (Figure \(3\)). Deforestation, cement manufacture, animal agriculture, the clearing of land, and the burning of forests are other human activities that release carbon dioxide. Methane (CH4) is produced when bacteria break down organic matter under anaerobic conditions. Anaerobic conditions can happen when organic matter is trapped underwater (such as in rice paddies) or in the intestines of herbivores. Methane can also be released from natural gas fields and the decomposition that occurs in landfills. Another source of methane is the melting of clathrates. Clathrates are frozen chunks of ice and methane found at the bottom of the ocean. When water warms, these chunks of ice melt and methane is released. As the ocean’s water temperature increases, the rate at which clathrates melt is increasing, releasing even more methane. This leads to increased levels of methane in the atmosphere, which further accelerates the rate of global warming. This is an example of the positive feedback loop that is leading to the rapid rate of increase of global temperatures.
Documented Results of Climate Change: Past and Present
Scientists have geological evidence of the consequences of long-ago climate change. Modern-day phenomena such as retreating glaciers and melting polar ice cause a continual rise in sea level. Meanwhile, changes in climate can negatively affect organisms.
Geological Climate Change
Global warming has been associated with at least one planet-wide extinction event during the geological past. The Permian extinction event occurred about 251 million years ago toward the end of the roughly 50-million-year-long geological time span known as the Permian period. This geologic time period was one of the three warmest periods in Earth’s geologic history. Scientists estimate that approximately 70 percent of the terrestrial plant and animal species and 84 percent of marine species became extinct, vanishing forever near the end of the Permian period. Organisms that had adapted to wet and warm climatic conditions, such as annual rainfall of 300–400 cm (118–157 in) and 20 °C–30 °C (68 °F–86 °F) in the tropical wet forest, may not have been able to survive the Permian climate change.
Link to Learning
Watch this NASA video to discover the mixed effects of global warming on plant growth. While scientists found that warmer temperatures in the 1980s and 1990s caused an increase in plant productivity, this advantage has since been counteracted by more frequent droughts.
Present Climate Change
A number of global events have occurred that may be attributed to climate change during our lifetimes. Glacier National Park in Montana is undergoing the retreat of many of its glaciers, a phenomenon known as glacier recession. In 1850, the area contained approximately 150 glaciers. By 2010, however, the park contained only about 24 glaciers greater than 25 acres in size. One of these glaciers is the Grinnell Glacier (Figure \(4\)) at Mount Gould. Between 1966 and 2005, the size of Grinnell Glacier shrank by 40 percent. Similarly, the mass of the ice sheets in Greenland and the Antarctic is decreasing: Greenland lost 150–250 km3 of ice per year between 2002 and 2006. In addition, the size and thickness of the Arctic sea ice is decreasing.
This loss of ice is leading to increases in the global sea level. On average, the sea is rising at a rate of 1.8 mm per year. However, between 1993 and 2010 the rate of sea level increase ranged between 2.9 and 3.4 mm per year. A variety of factors affect the volume of water in the ocean, including the temperature of the water (the density of water is related to its temperature) and the amount of water found in rivers, lakes, glaciers, polar ice caps, and sea ice. As glaciers and polar ice caps melt, there is a significant contribution of liquid water that was previously frozen.
In addition to some abiotic conditions changing in response to climate change, many organisms are also being affected by the changes in temperature. Temperature and precipitation play key roles in determining the geographic distribution and phenology of plants and animals. (Phenology is the study of the effects of climatic conditions on the timing of periodic lifecycle events, such as flowering in plants or migration in birds.) Researchers have shown that 385 plant species in Great Britain are flowering 4.5 days sooner than was recorded earlier during the previous 40 years. In addition, insect-pollinated species were more likely to flower earlier than wind-pollinated species. The impact of changes in flowering date would be mitigated if the insect pollinators emerged earlier. This mismatched timing of plants and pollinators could result in injurious ecosystem effects because, for continued survival, insect-pollinated plants must flower when their pollinators are present.
Summary
The Earth has gone through periodic cycles of increases and decreases in temperature. During the past 2000 years, the Medieval Climate Anomaly was a warmer period, while the Little Ice Age was unusually cool. Both of these irregularities can be explained by natural causes of changes in climate, and, although the temperature changes were small, they had significant effects. Natural drivers of climate change include Milankovitch cycles, changes in solar activity, and volcanic eruptions. None of these factors, however, leads to rapid increases in global temperature or sustained increases in carbon dioxide. The burning of fossil fuels is an important source of greenhouse gases, which plays a major role in the greenhouse effect. Long ago, global warming resulted in the Permian extinction: a large-scale extinction event that is documented in the fossil record. Currently, modern-day climate change is associated with the increased melting of glaciers and polar ice sheets, resulting in a gradual increase in sea level. Plants and animals can also be affected by global climate change when the timing of seasonal events, such as flowering or pollination, is affected by global warming.
Glossary
clathrates
frozen chunks of ice and methane found at the bottom of the ocean
climate
long-term, predictable atmospheric conditions present in a specific area
global climate change
altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide
greenhouse effect
warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere
greenhouse gases
atmospheric gases such as carbon dioxide and methane that absorb and emit radiation, thus trapping heat in Earth’s atmosphere
haze-effect cooling
effect of the gases and solids from a volcanic eruption on global climate
Milankovitch cycles
cyclic changes in the Earth's orbit that may affect climate
solar intensity
amount of solar power energy the sun emits in a given amount of time
weather
conditions of the atmosphere during a short period of time | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.05%3A_Human_Impacts_of_the_Biosphere-_Pollution_and_Resource_Depletion/57.5.01%3A_Climate_and_the_Effects_of_Global.txt |
Skills to Develop
• Discuss the biogeochemical cycles of water, carbon, nitrogen, phosphorus, and sulfur
• Explain how human activities have impacted these cycles and the potential consequences for Earth
Energy flows directionally through ecosystems, entering as sunlight (or inorganic molecules for chemoautotrophs) and leaving as heat during the many transfers between trophic levels. However, the matter that makes up living organisms is conserved and recycled. The six most common elements associated with organic molecules—carbon, nitrogen, hydrogen, oxygen, phosphorus, and sulfur—take a variety of chemical forms and may exist for long periods in the atmosphere, on land, in water, or beneath the Earth’s surface. Geologic processes, such as weathering, erosion, water drainage, and the subduction of the continental plates, all play a role in this recycling of materials. Because geology and chemistry have major roles in the study of this process, the recycling of inorganic matter between living organisms and their environment is called a biogeochemical cycle.
Water contains hydrogen and oxygen, which is essential to all living processes. The hydrosphere is the area of the Earth where water movement and storage occurs: as liquid water on the surface and beneath the surface or frozen (rivers, lakes, oceans, groundwater, polar ice caps, and glaciers), and as water vapor in the atmosphere. Carbon is found in all organic macromolecules and is an important constituent of fossil fuels. Nitrogen is a major component of our nucleic acids and proteins and is critical to human agriculture. Phosphorus, a major component of nucleic acid (along with nitrogen), is one of the main ingredients in artificial fertilizers used in agriculture and their associated environmental impacts on our surface water. Sulfur, critical to the 3–D folding of proteins (as in disulfide binding), is released into the atmosphere by the burning of fossil fuels, such as coal.
The cycling of these elements is interconnected. For example, the movement of water is critical for the leaching of nitrogen and phosphate into rivers, lakes, and oceans. Furthermore, the ocean itself is a major reservoir for carbon. Thus, mineral nutrients are cycled, either rapidly or slowly, through the entire biosphere, from one living organism to another, and between the biotic and abiotic world.
The Water (Hydrologic) Cycle
Water is the basis of all living processes. The human body is more than 1/2 water and human cells are more than 70 percent water. Thus, most land animals need a supply of fresh water to survive. However, when examining the stores of water on Earth, 97.5 percent of it is non-potable salt water (Figure $1$). Of the remaining water, 99 percent is locked underground as water or as ice. Thus, less than 1 percent of fresh water is easily accessible from lakes and rivers. Many living things, such as plants, animals, and fungi, are dependent on the small amount of fresh surface water supply, a lack of which can have massive effects on ecosystem dynamics. Humans, of course, have developed technologies to increase water availability, such as digging wells to harvest groundwater, storing rainwater, and using desalination to obtain drinkable water from the ocean. Although this pursuit of drinkable water has been ongoing throughout human history, the supply of fresh water is still a major issue in modern times.
Water cycling is extremely important to ecosystem dynamics. Water has a major influence on climate and, thus, on the environments of ecosystems, some located on distant parts of the Earth. Most of the water on Earth is stored for long periods in the oceans, underground, and as ice. Figure $2$ illustrates the average time that an individual water molecule may spend in the Earth’s major water reservoirs. Residence time is a measure of the average time an individual water molecule stays in a particular reservoir. A large amount of the Earth’s water is locked in place in these reservoirs as ice, beneath the ground, and in the ocean, and, thus, is unavailable for short-term cycling (only surface water can evaporate).
There are various processes that occur during the cycling of water, shown in Figure $3$. These processes include the following:
• evaporation/sublimation
• condensation/precipitation
• subsurface water flow
• surface runoff/snowmelt
• streamflow
The water cycle is driven by the sun’s energy as it warms the oceans and other surface waters. This leads to the evaporation (water to water vapor) of liquid surface water and the sublimation (ice to water vapor) of frozen water, which deposits large amounts of water vapor into the atmosphere. Over time, this water vapor condenses into clouds as liquid or frozen droplets and is eventually followed by precipitation (rain or snow), which returns water to the Earth’s surface. Rain eventually permeates into the ground, where it may evaporate again if it is near the surface, flow beneath the surface, or be stored for long periods. More easily observed is surface runoff: the flow of fresh water either from rain or melting ice. Runoff can then make its way through streams and lakes to the oceans or flow directly to the oceans themselves.
Link to Learning
Head to this website to learn more about the world’s fresh water supply.
Rain and surface runoff are major ways in which minerals, including carbon, nitrogen, phosphorus, and sulfur, are cycled from land to water. The environmental effects of runoff will be discussed later as these cycles are described.
The Carbon Cycle
Carbon is the second most abundant element in living organisms. Carbon is present in all organic molecules, and its role in the structure of macromolecules is of primary importance to living organisms. Carbon compounds contain especially high energy, particularly those derived from fossilized organisms, mainly plants, which humans use as fuel. Since the 1800s, the number of countries using massive amounts of fossil fuels has increased. Since the beginning of the Industrial Revolution, global demand for the Earth’s limited fossil fuel supplies has risen; therefore, the amount of carbon dioxide in our atmosphere has increased. This increase in carbon dioxide has been associated with climate change and other disturbances of the Earth’s ecosystems and is a major environmental concern worldwide. Thus, the “carbon footprint” is based on how much carbon dioxide is produced and how much fossil fuel countries consume.
The carbon cycle is most easily studied as two interconnected sub-cycles: one dealing with rapid carbon exchange among living organisms and the other dealing with the long-term cycling of carbon through geologic processes. The entire carbon cycle is shown in Figure $4$.
Link to Learning
Click this link to read information about the United States Carbon Cycle Science Program.
The Biological Carbon Cycle
Living organisms are connected in many ways, even between ecosystems. A good example of this connection is the exchange of carbon between autotrophs and heterotrophs within and between ecosystems by way of atmospheric carbon dioxide. Carbon dioxide is the basic building block that most autotrophs use to build multi-carbon, high energy compounds, such as glucose. The energy harnessed from the sun is used by these organisms to form the covalent bonds that link carbon atoms together. These chemical bonds thereby store this energy for later use in the process of respiration. Most terrestrial autotrophs obtain their carbon dioxide directly from the atmosphere, while marine autotrophs acquire it in the dissolved form (carbonic acid, H2CO3). However carbon dioxide is acquired, a by-product of the process is oxygen. The photosynthetic organisms are responsible for depositing approximately 21 percent oxygen content of the atmosphere that we observe today.
Heterotrophs and autotrophs are partners in biological carbon exchange (especially the primary consumers, largely herbivores). Heterotrophs acquire the high-energy carbon compounds from the autotrophs by consuming them, and breaking them down by respiration to obtain cellular energy, such as ATP. The most efficient type of respiration, aerobic respiration, requires oxygen obtained from the atmosphere or dissolved in water. Thus, there is a constant exchange of oxygen and carbon dioxide between the autotrophs (which need the carbon) and the heterotrophs (which need the oxygen). Gas exchange through the atmosphere and water is one way that the carbon cycle connects all living organisms on Earth.
The Biogeochemical Carbon Cycle
The movement of carbon through the land, water, and air is complex, and in many cases, it occurs much more slowly geologically than as seen between living organisms. Carbon is stored for long periods in what are known as carbon reservoirs, which include the atmosphere, bodies of liquid water (mostly oceans), ocean sediment, soil, land sediments (including fossil fuels), and the Earth’s interior.
As stated, the atmosphere is a major reservoir of carbon in the form of carbon dioxide and is essential to the process of photosynthesis. The level of carbon dioxide in the atmosphere is greatly influenced by the reservoir of carbon in the oceans. The exchange of carbon between the atmosphere and water reservoirs influences how much carbon is found in each location, and each one affects the other reciprocally. Carbon dioxide (CO2) from the atmosphere dissolves in water and combines with water molecules to form carbonic acid, and then it ionizes to carbonate and bicarbonate ions (Figure $5$)
The equilibrium coefficients are such that more than 90 percent of the carbon in the ocean is found as bicarbonate ions. Some of these ions combine with seawater calcium to form calcium carbonate (CaCO3), a major component of marine organism shells. These organisms eventually form sediments on the ocean floor. Over geologic time, the calcium carbonate forms limestone, which comprises the largest carbon reservoir on Earth.
On land, carbon is stored in soil as a result of the decomposition of living organisms (by decomposers) or from weathering of terrestrial rock and minerals. This carbon can be leached into the water reservoirs by surface runoff. Deeper underground, on land and at sea, are fossil fuels: the anaerobically decomposed remains of plants that take millions of years to form. Fossil fuels are considered a non-renewable resource because their use far exceeds their rate of formation. A non-renewable resource, such as fossil fuel, is either regenerated very slowly or not at all. Another way for carbon to enter the atmosphere is from land (including land beneath the surface of the ocean) by the eruption of volcanoes and other geothermal systems. Carbon sediments from the ocean floor are taken deep within the Earth by the process of subduction: the movement of one tectonic plate beneath another. Carbon is released as carbon dioxide when a volcano erupts or from volcanic hydrothermal vents.
Carbon dioxide is also added to the atmosphere by the animal husbandry practices of humans. The large numbers of land animals raised to feed the Earth’s growing population results in increased carbon dioxide levels in the atmosphere due to farming practices and the respiration and methane production. This is another example of how human activity indirectly affects biogeochemical cycles in a significant way. Although much of the debate about the future effects of increasing atmospheric carbon on climate change focuses on fossils fuels, scientists take natural processes, such as volcanoes and respiration, into account as they model and predict the future impact of this increase.
The Nitrogen Cycle
Getting nitrogen into the living world is difficult. Plants and phytoplankton are not equipped to incorporate nitrogen from the atmosphere (which exists as tightly bonded, triple covalent N2) even though this molecule comprises approximately 78 percent of the atmosphere. Nitrogen enters the living world via free-living and symbiotic bacteria, which incorporate nitrogen into their macromolecules through nitrogen fixation (conversion of N2). Cyanobacteria live in most aquatic ecosystems where sunlight is present; they play a key role in nitrogen fixation. Cyanobacteria are able to use inorganic sources of nitrogen to “fix” nitrogen. Rhizobium bacteria live symbiotically in the root nodules of legumes (such as peas, beans, and peanuts) and provide them with the organic nitrogen they need. Free-living bacteria, such as Azotobacter, are also important nitrogen fixers.
Organic nitrogen is especially important to the study of ecosystem dynamics since many ecosystem processes, such as primary production and decomposition, are limited by the available supply of nitrogen. As shown in Figure $6$, the nitrogen that enters living systems by nitrogen fixation is successively converted from organic nitrogen back into nitrogen gas by bacteria. This process occurs in three steps in terrestrial systems: ammonification, nitrification, and denitrification. First, the ammonification process converts nitrogenous waste from living animals or from the remains of dead animals into ammonium (NH4+) by certain bacteria and fungi. Second, the ammonium is converted to nitrites (NO2) by nitrifying bacteria, such as Nitrosomonas, through nitrification. Subsequently, nitrites are converted to nitrates (NO3) by similar organisms. Third, the process of denitrification occurs, whereby bacteria, such as Pseudomonas and Clostridium, convert the nitrates into nitrogen gas, allowing it to re-enter the atmosphere.
Exercise
Which of the following statements about the nitrogen cycle is false?
1. Ammonification converts organic nitrogenous matter from living organisms into ammonium (NH4+).
2. Denitrification by bacteria converts nitrates (NO3) to nitrogen gas (N2).
3. Nitrification by bacteria converts nitrates (NO3) to nitrites (NO2).
4. Nitrogen fixing bacteria convert nitrogen gas (N2) into organic compounds.
Answer
C: Nitrification by bacteria converts nitrates (NO3) to nitrites (NO2).
Human activity can release nitrogen into the environment by two primary means: the combustion of fossil fuels, which releases different nitrogen oxides, and by the use of artificial fertilizers in agriculture, which are then washed into lakes, streams, and rivers by surface runoff. Atmospheric nitrogen is associated with several effects on Earth’s ecosystems including the production of acid rain (as nitric acid, HNO3) and greenhouse gas (as nitrous oxide, N2O) potentially causing climate change. A major effect from fertilizer runoff is saltwater and freshwater eutrophication, a process whereby nutrient runoff causes the excess growth of microorganisms, depleting dissolved oxygen levels and killing ecosystem fauna.
A similar process occurs in the marine nitrogen cycle, where the ammonification, nitrification, and denitrification processes are performed by marine bacteria. Some of this nitrogen falls to the ocean floor as sediment, which can then be moved to land in geologic time by uplift of the Earth’s surface and thereby incorporated into terrestrial rock. Although the movement of nitrogen from rock directly into living systems has been traditionally seen as insignificant compared with nitrogen fixed from the atmosphere, a recent study showed that this process may indeed be significant and should be included in any study of the global nitrogen cycle.1
The Phosphorus Cycle
Phosphorus is an essential nutrient for living processes; it is a major component of nucleic acid and phospholipids, and, as calcium phosphate, makes up the supportive components of our bones. Phosphorus is often the limiting nutrient (necessary for growth) in aquatic ecosystems (Figure $7$).
Phosphorus occurs in nature as the phosphate ion (PO43−). In addition to phosphate runoff as a result of human activity, natural surface runoff occurs when it is leached from phosphate-containing rock by weathering, thus sending phosphates into rivers, lakes, and the ocean. This rock has its origins in the ocean. Phosphate-containing ocean sediments form primarily from the bodies of ocean organisms and from their excretions. However, in remote regions, volcanic ash, aerosols, and mineral dust may also be significant phosphate sources. This sediment then is moved to land over geologic time by the uplifting of areas of the Earth’s surface.
Phosphorus is also reciprocally exchanged between phosphate dissolved in the ocean and marine ecosystems. The movement of phosphate from the ocean to the land and through the soil is extremely slow, with the average phosphate ion having an oceanic residence time between 20,000 and 100,000 years.
Excess phosphorus and nitrogen that enters these ecosystems from fertilizer runoff and from sewage causes excessive growth of microorganisms and depletes the dissolved oxygen, which leads to the death of many ecosystem fauna, such as shellfish and finfish. This process is responsible for dead zones in lakes and at the mouths of many major rivers (Figure $8$).
A dead zone is an area within a freshwater or marine ecosystem where large areas are depleted of their normal flora and fauna; these zones can be caused by eutrophication, oil spills, dumping of toxic chemicals, and other human activities. The number of dead zones has been increasing for several years, and more than 400 of these zones were present as of 2008. One of the worst dead zones is off the coast of the United States in the Gulf of Mexico, where fertilizer runoff from the Mississippi River basin has created a dead zone of over 8463 square miles. Phosphate and nitrate runoff from fertilizers also negatively affect several lake and bay ecosystems including the Chesapeake Bay in the eastern United States.
Everyday Connection: Chesapeake Bay
The Chesapeake Bay has long been valued as one of the most scenic areas on Earth; it is now in distress and is recognized as a declining ecosystem. In the 1970s, the Chesapeake Bay was one of the first ecosystems to have identified dead zones, which continue to kill many fish and bottom-dwelling species, such as clams, oysters, and worms. Several species have declined in the Chesapeake Bay due to surface water runoff containing excess nutrients from artificial fertilizer used on land. The source of the fertilizers (with high nitrogen and phosphate content) is not limited to agricultural practices. There are many nearby urban areas and more than 150 rivers and streams empty into the bay that are carrying fertilizer runoff from lawns and gardens. Thus, the decline of the Chesapeake Bay is a complex issue and requires the cooperation of industry, agriculture, and everyday homeowners.
Of particular interest to conservationists is the oyster population; it is estimated that more than 200,000 acres of oyster reefs existed in the bay in the 1700s, but that number has now declined to only 36,000 acres. Oyster harvesting was once a major industry for Chesapeake Bay, but it declined 88 percent between 1982 and 2007. This decline was due not only to fertilizer runoff and dead zones but also to overharvesting. Oysters require a certain minimum population density because they must be in close proximity to reproduce. Human activity has altered the oyster population and locations, greatly disrupting the ecosystem.
The restoration of the oyster population in the Chesapeake Bay has been ongoing for several years with mixed success. Not only do many people find oysters good to eat, but they also clean up the bay. Oysters are filter feeders, and as they eat, they clean the water around them. In the 1700s, it was estimated that it took only a few days for the oyster population to filter the entire volume of the bay. Today, with changed water conditions, it is estimated that the present population would take nearly a year to do the same job.
Restoration efforts have been ongoing for several years by non-profit organizations, such as the Chesapeake Bay Foundation. The restoration goal is to find a way to increase population density so the oysters can reproduce more efficiently. Many disease-resistant varieties (developed at the Virginia Institute of Marine Science for the College of William and Mary) are now available and have been used in the construction of experimental oyster reefs. Efforts to clean and restore the bay by Virginia and Delaware have been hampered because much of the pollution entering the bay comes from other states, which stresses the need for inter-state cooperation to gain successful restoration.
The new, hearty oyster strains have also spawned a new and economically viable industry—oyster aquaculture—which not only supplies oysters for food and profit, but also has the added benefit of cleaning the bay.
The Sulfur Cycle
Sulfur is an essential element for the macromolecules of living things. As a part of the amino acid cysteine, it is involved in the formation of disulfide bonds within proteins, which help to determine their 3-D folding patterns, and hence their functions. As shown in Figure $10$, sulfur cycles between the oceans, land, and atmosphere. Atmospheric sulfur is found in the form of sulfur dioxide (SO2) and enters the atmosphere in three ways: from the decomposition of organic molecules, from volcanic activity and geothermal vents, and from the burning of fossil fuels by humans.
On land, sulfur is deposited in four major ways: precipitation, direct fallout from the atmosphere, rock weathering, and geothermal vents (Figure $11$). Atmospheric sulfur is found in the form of sulfur dioxide (SO2), and as rain falls through the atmosphere, sulfur is dissolved in the form of weak sulfuric acid (H2SO4). Sulfur can also fall directly from the atmosphere in a process called fallout. Also, the weathering of sulfur-containing rocks releases sulfur into the soil. These rocks originate from ocean sediments that are moved to land by the geologic uplifting of ocean sediments. Terrestrial ecosystems can then make use of these soil sulfates ($\text{SO}_4^{2-}$), and upon the death and decomposition of these organisms, release the sulfur back into the atmosphere as hydrogen sulfide (H2S) gas.
Sulfur enters the ocean via runoff from land, from atmospheric fallout, and from underwater geothermal vents. Some ecosystems rely on chemoautotrophs using sulfur as a biological energy source. This sulfur then supports marine ecosystems in the form of sulfates.
Human activities have played a major role in altering the balance of the global sulfur cycle. The burning of large quantities of fossil fuels, especially from coal, releases larger amounts of hydrogen sulfide gas into the atmosphere. As rain falls through this gas, it creates the phenomenon known as acid rain. Acid rain is corrosive rain caused by rainwater falling to the ground through sulfur dioxide gas, turning it into weak sulfuric acid, which causes damage to aquatic ecosystems. Acid rain damages the natural environment by lowering the pH of lakes, which kills many of the resident fauna; it also affects the man-made environment through the chemical degradation of buildings. For example, many marble monuments, such as the Lincoln Memorial in Washington, DC, have suffered significant damage from acid rain over the years. These examples show the wide-ranging effects of human activities on our environment and the challenges that remain for our future.
Link to Learning
Click this link to learn more about global climate change.
Summary
Mineral nutrients are cycled through ecosystems and their environment. Of particular importance are water, carbon, nitrogen, phosphorus, and sulfur. All of these cycles have major impacts on ecosystem structure and function. As human activities have caused major disturbances to these cycles, their study and modeling is especially important. A variety of human activities, such as pollution, oil spills, and other events have damaged ecosystems, potentially causing global climate change. The health of Earth depends on understanding these cycles and how to protect the environment from irreversible damage.
Footnotes
1. 1 Scott L. Morford, Benjamin Z. Houlton, and Randy A. Dahlgren, “Increased Forest Ecosystem Carbon and Nitrogen Storage from Nitrogen Rich Bedrock,” Nature 477, no. 7362 (2011): 78–81.
Glossary
acid rain
corrosive rain caused by rainwater falling to the ground through sulfur dioxide gas, turning it into weak sulfuric acid; can damage structures and ecosystems
biogeochemical cycle
cycling of mineral nutrients through ecosystems and through the non-living world
dead zone
area within an ecosystem in lakes and near the mouths of rivers where large areas of ecosystems are depleted of their normal flora and fauna; these zones can be caused by eutrophication, oil spills, dumping of toxic chemicals, and other human activities
eutrophication
process whereby nutrient runoff causes the excess growth of microorganisms, depleting dissolved oxygen levels and killing ecosystem fauna
fallout
direct deposit of solid minerals on land or in the ocean from the atmosphere
hydrosphere
area of the Earth where water movement and storage occurs
non-renewable resource
resource, such as fossil fuel, that is either regenerated very slowly or not at all
residence time
measure of the average time an individual water molecule stays in a particular reservoir
subduction
movement of one tectonic plate beneath another | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.05%3A_Human_Impacts_of_the_Biosphere-_Pollution_and_Resource_Depletion/57.5.02%3A_Biogeochemical_Cycles.txt |
Skills to Develop
• Identify significant threats to biodiversity
• Explain the effects of habitat loss, exotic species, and hunting on biodiversity
• Identify the early and predicted effects of climate change on biodiversity
The core threat to biodiversity on the planet, and therefore a threat to human welfare, is the combination of human population growth and resource exploitation. The human population requires resources to survive and grow, and those resources are being removed unsustainably from the environment. The three greatest proximate threats to biodiversity are habitat loss, overharvesting, and introduction of exotic species. The first two of these are a direct result of human population growth and resource use. The third results from increased mobility and trade. A fourth major cause of extinction, anthropogenic climate change, has not yet had a large impact, but it is predicted to become significant during this century. Global climate change is also a consequence of human population needs for energy and the use of fossil fuels to meet those needs (Figure \(1\)). Environmental issues, such as toxic pollution, have specific targeted effects on species, but they are not generally seen as threats at the magnitude of the others.
Habitat Loss
Humans rely on technology to modify their environment and replace certain functions that were once performed by the natural ecosystem. Other species cannot do this. Elimination of their ecosystem—whether it is a forest, a desert, a grassland, a freshwater estuarine, or a marine environment—will kill the individuals in the species. Remove the entire habitat within the range of a species and, unless they are one of the few species that do well in human-built environments, the species will become extinct. Human destruction of habitats accelerated in the latter half of the twentieth century. Consider the exceptional biodiversity of Sumatra: it is home to one species of orangutan, a species of critically endangered elephant, and the Sumatran tiger, but half of Sumatra’s forest is now gone. The neighboring island of Borneo, home to the other species of orangutan, has lost a similar area of forest. Forest loss continues in protected areas of Borneo. The orangutan in Borneo is listed as endangered by the International Union for Conservation of Nature (IUCN), but it is simply the most visible of thousands of species that will not survive the disappearance of the forests of Borneo. The forests are removed for timber and to plant palm oil plantations (Figure \(2\)). Palm oil is used in many products including food products, cosmetics, and biodiesel in Europe. A five-year estimate of global forest cover loss for the years 2000–2005 was 3.1 percent. In the humid tropics where forest loss is primarily from timber extraction, 272,000 km2 was lost out of a global total of 11,564,000 km2 (or 2.4 percent). In the tropics, these losses certainly also represent the extinction of species because of high levels of endemism.
Everyday Connection: Preventing Habitat Destruction with Wise Wood Choices
Most consumers do not imagine that the home improvement products they buy might be contributing to habitat loss and species extinctions. Yet the market for illegally harvested tropical timber is huge, and the wood products often find themselves in building supply stores in the United States. One estimate is that 10 percent of the imported timber stream in the United States, which is the world’s largest consumer of wood products, is potentially illegally logged. In 2006, this amounted to \$3.6 billion in wood products. Most of the illegal products are imported from countries that act as intermediaries and are not the originators of the wood.
How is it possible to determine if a wood product, such as flooring, was harvested sustainably or even legally? The Forest Stewardship Council (FSC) certifies sustainably harvested forest products, therefore, looking for their certification on flooring and other hardwood products is one way to ensure that the wood has not been taken illegally from a tropical forest. Certification applies to specific products, not to a producer; some producers’ products may not have certification while other products are certified. While there are other industry-backed certifications other than the FSC, these are unreliable due to lack of independence from the industry. Another approach is to buy domestic wood species. While it would be great if there was a list of legal versus illegal wood products, it is not that simple. Logging and forest management laws vary from country to country; what is illegal in one country may be legal in another. Where and how a product is harvested and whether the forest from which it comes is being maintained sustainably all factor into whether a wood product will be certified by the FSC. It is always a good idea to ask questions about where a wood product came from and how the supplier knows that it was harvested legally.
Habitat destruction can affect ecosystems other than forests. Rivers and streams are important ecosystems and are frequently modified through land development and from damming or water removal. Damming of rivers affects the water flow and access to all parts of a river. Differing flow regimes can reduce or eliminate populations that are adapted to these changes in flow patterns. For example, an estimated 91percent of river lengths in the United States have been developed: they have modifications like dams, to create energy or store water; levees, to prevent flooding; or dredging or rerouting, to create land that is more suitable for human development. Many fish species in the United States, especially rare species or species with restricted distributions, have seen declines caused by river damming and habitat loss. Research has confirmed that species of amphibians that must carry out parts of their life cycles in both aquatic and terrestrial habitats have a greater chance of suffering population declines and extinction because of the increased likelihood that one of their habitats or access between them will be lost.
Overharvesting
Overharvesting is a serious threat to many species, but particularly to aquatic species. There are many examples of regulated commercial fisheries monitored by fisheries scientists that have nevertheless collapsed. The western Atlantic cod fishery is the most spectacular recent collapse. While it was a hugely productive fishery for 400 years, the introduction of modern factory trawlers in the 1980s and the pressure on the fishery led to it becoming unsustainable. The causes of fishery collapse are both economic and political in nature. Most fisheries are managed as a common (shared) resource even when the fishing territory lies within a country’s territorial waters. Common resources are subject to an economic pressure known as the tragedy of the commons in which essentially no fisher has a motivation to exercise restraint in harvesting a fishery when it is not owned by that fisher. The natural outcome of harvests of resources held in common is their overexploitation. While large fisheries are regulated to attempt to avoid this pressure, it still exists in the background. This overexploitation is exacerbated when access to the fishery is open and unregulated and when technology gives fishers the ability to overfish. In a few fisheries, the biological growth of the resource is less than the potential growth of the profits made from fishing if that time and money were invested elsewhere. In these cases—whales are an example—economic forces will always drive toward fishing the population to extinction.
Link to Learning
Explore a U.S. Fish & Wildlife Service interactive map of critical habitat for endangered and threatened species in the United States. To begin, select “Visit the online mapper.”
For the most part, fishery extinction is not equivalent to biological extinction—the last fish of a species is rarely fished out of the ocean. At the same time, fishery extinction is still harmful to fish species and their ecosystems. There are some instances in which true extinction is a possibility. Whales have slow-growing populations and are at risk of complete extinction through hunting. There are some species of sharks with restricted distributions that are at risk of extinction. The groupers are another population of generally slow-growing fishes that, in the Caribbean, includes a number of species that are at risk of extinction from overfishing.
Coral reefs are extremely diverse marine ecosystems that face peril from several processes. Reefs are home to 1/3 of the world’s marine fish species—about 4,000 species—despite making up only 1 percent of marine habitat. Most home marine aquaria are stocked with wild-caught organisms, not cultured organisms. Although no species is known to have been driven extinct by the pet trade in marine species, there are studies showing that populations of some species have declined in response to harvesting, indicating that the harvest is not sustainable at those levels. There are concerns about the effect of the pet trade on some terrestrial species such as turtles, amphibians, birds, plants, and even the orangutan.
Bush meat is the generic term used for wild animals killed for food. Hunting is practiced throughout the world, but hunting practices, particularly in equatorial Africa and parts of Asia, are believed to threaten several species with extinction. Traditionally, bush meat in Africa was hunted to feed families directly; however, recent commercialization of the practice now has bush meat available in grocery stores, which has increased harvest rates to the level of unsustainability. Additionally, human population growth has increased the need for protein foods that are not being met from agriculture. Species threatened by the bush meat trade are mostly mammals including many primates living in the Congo basin.
Exotic Species
Exotic species are species that have been intentionally or unintentionally introduced by humans into an ecosystem in which they did not evolve. Such introductions likely occur frequently as natural phenomena. For example, Kudzu (Pueraria lobata), which is native to Japan, was introduced in the United States in 1876. It was later planted for soil conservation. Problematically, it grows too well in the southeastern United States—up to a foot a day. It is now a pest species and covers over 7 million acres in the southeastern United States. If an introduced species is able to survive in its new habitat, that introduction is now reflected in the observed range of the species. Human transportation of people and goods, including the intentional transport of organisms for trade, has dramatically increased the introduction of species into new ecosystems, sometimes at distances that are well beyond the capacity of the species to ever travel itself and outside the range of the species’ natural predators.
Most exotic species introductions probably fail because of the low number of individuals introduced or poor adaptation to the ecosystem they enter. Some species, however, possess preadaptations that can make them especially successful in a new ecosystem. These exotic species often undergo dramatic population increases in their new habitat and reset the ecological conditions in the new environment, threatening the species that exist there. For this reason, exotic species are also called invasive species. Exotic species can threaten other species through competition for resources, predation, or disease.
Link to Learning
Explore an interactive global database of exotic or invasive species.
Lakes and islands are particularly vulnerable to extinction threats from introduced species. In Lake Victoria, as mentioned earlier, the intentional introduction of the Nile perch was largely responsible for the extinction of about 200 species of cichlids. The accidental introduction of the brown tree snake via aircraft (Figure \(3\)) from the Solomon Islands to Guam in 1950 has led to the extinction of three species of birds and three to five species of reptiles endemic to the island. Several other species are still threatened. The brown tree snake is adept at exploiting human transportation as a means to migrate; one was even found on an aircraft arriving in Corpus Christi, Texas. Constant vigilance on the part of airport, military, and commercial aircraft personnel is required to prevent the snake from moving from Guam to other islands in the Pacific, especially Hawaii. Islands do not make up a large area of land on the globe, but they do contain a disproportionate number of endemic species because of their isolation from mainland ancestors.
It now appears that the global decline in amphibian species recognized in the 1990s is, in some part, caused by the fungus Batrachochytrium dendrobatidis, which causes the disease chytridiomycosis (Figure \(4\)). There is evidence that the fungus is native to Africa and may have been spread throughout the world by transport of a commonly used laboratory and pet species: the African clawed toad (Xenopus laevis). It may well be that biologists themselves are responsible for spreading this disease worldwide. The North American bullfrog, Rana catesbeiana, which has also been widely introduced as a food animal but which easily escapes captivity, survives most infections of Batrachochytrium dendrobatidis and can act as a reservoir for the disease.
Early evidence suggests that another fungal pathogen, Geomyces destructans, introduced from Europe is responsible for white-nose syndrome, which infects cave-hibernating bats in eastern North America and has spread from a point of origin in western New York State (Figure \(5\)). The disease has decimated bat populations and threatens extinction of species already listed as endangered: the Indiana bat, Myotis sodalis, and potentially the Virginia big-eared bat, Corynorhinus townsendii virginianus. How the fungus was introduced is unclear, but one logical presumption would be that recreational cavers unintentionally brought the fungus on clothes or equipment from Europe.
Climate Change
Climate change, and specifically the anthropogenic (meaning, caused by humans) warming trend presently underway, is recognized as a major extinction threat, particularly when combined with other threats such as habitat loss. Scientists disagree about the likely magnitude of the effects, with extinction rate estimates ranging from 15 percent to 40 percent of species committed to extinction by 2050. Scientists do agree, however, that climate change will alter regional climates, including rainfall and snowfall patterns, making habitats less hospitable to the species living in them. The warming trend will shift colder climates toward the north and south poles, forcing species to move with their adapted climate norms while facing habitat gaps along the way. The shifting ranges will impose new competitive regimes on species as they find themselves in contact with other species not present in their historic range. One such unexpected species contact is between polar bears and grizzly bears. Previously, these two species had separate ranges. Now, their ranges are overlapping and there are documented cases of these two species mating and producing viable offspring. Changing climates also throw off species’ delicate timing adaptations to seasonal food resources and breeding times. Many contemporary mismatches to shifts in resource availability and timing have already been documented.
Range shifts are already being observed: for example, some European bird species ranges have moved 91 km northward. The same study suggested that the optimal shift based on warming trends was double that distance, suggesting that the populations are not moving quickly enough. Range shifts have also been observed in plants, butterflies, other insects, freshwater fishes, reptiles, and mammals.
Climate gradients will also move up mountains, eventually crowding species higher in altitude and eliminating the habitat for those species adapted to the highest elevations. Some climates will completely disappear. The rate of warming appears to be accelerated in the arctic, which is recognized as a serious threat to polar bear populations that require sea ice to hunt seals during the winter months: seals are the only source of protein available to polar bears. A trend to decreasing sea ice coverage has occurred since observations began in the mid-twentieth century. The rate of decline observed in recent years is far greater than previously predicted by climate models.
Finally, global warming will raise ocean levels due to melt water from glaciers and the greater volume of warmer water. Shorelines will be inundated, reducing island size, which will have an effect on some species, and a number of islands will disappear entirely. Additionally, the gradual melting and subsequent refreezing of the poles, glaciers, and higher elevation mountains—a cycle that has provided freshwater to environments for centuries—will also be jeopardized. This could result in an overabundance of salt water and a shortage of fresh water.
Summary
The core threats to biodiversity are human population growth and unsustainable resource use. To date, the most significant causes of extinctions are habitat loss, introduction of exotic species, and overharvesting. Climate change is predicted to be a significant cause of extinctions in the coming century. Habitat loss occurs through deforestation, damming of rivers, and other activities. Overharvesting is a threat particularly to aquatic species, while the taking of bush meat in the humid tropics threatens many species in Asia, Africa, and the Americas. Exotic species have been the cause of a number of extinctions and are especially damaging to islands and lakes. Exotic species’ introductions are increasing because of the increased mobility of human populations and growing global trade and transportation. Climate change is forcing range changes that may lead to extinction. It is also affecting adaptations to the timing of resource availability that negatively affects species in seasonal environments. The impacts of climate change are greatest in the arctic. Global warming will also raise sea levels, eliminating some islands and reducing the area of all others.
Art Connections
Converting a prairie to a farm field is an example of ________.
1. overharvesting
2. habitat loss
3. exotic species
4. climate change
Answer
B
Glossary
bush meat
wild-caught animal used as food (typically mammals, birds, and reptiles); usually referring to hunting in the tropics of sub-Saharan Africa, Asia, and the Americas
chytridiomycosis
disease of amphibians caused by the fungus Batrachochytrium dendrobatidis; thought to be a major cause of the global amphibian decline
exotic species
(also, invasive species) species that has been introduced to an ecosystem in which it did not evolve
tragedy of the commons
economic principle that resources held in common will inevitably be overexploited
white-nose syndrome
disease of cave-hibernating bats in the eastern United States and Canada associated with the fungus Geomyces destructans | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.05%3A_Human_Impacts_of_the_Biosphere-_Pollution_and_Resource_Depletion/57.5.03%3A_Threats_to_Biodiversity.txt |
Skills to Develop
• Define global climate change
• Summarize the effects of the Industrial Revolution on global atmospheric carbon dioxide concentration
• Describe three natural factors affecting long-term global climate
• List two or more greenhouse gases and describe their role in the greenhouse effect
All biomes are universally affected by global conditions, such as climate, that ultimately shape each biome’s environment. Scientists who study climate have noted a series of marked changes that have gradually become increasingly evident during the last sixty years. Global climate change is the term used to describe altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide.
Climate and Weather
A common misconception about global climate change is that a specific weather event occurring in a particular region (for example, a very cool week in June in central Indiana) is evidence of global climate change. However, a cold week in June is a weather-related event and not a climate-related one. These misconceptions often arise because of confusion over the terms climate and weather.
Climate refers to the long-term, predictable atmospheric conditions of a specific area. The climate of a biome is characterized by having consistent temperature and annual rainfall ranges. Climate does not address the amount of rain that fell on one particular day in a biome or the colder-than-average temperatures that occurred on one day. In contrast, weather refers to the conditions of the atmosphere during a short period of time. Weather forecasts are usually made for 48-hour cycles. Long-range weather forecasts are available but can be unreliable.
To better understand the difference between climate and weather, imagine that you are planning an outdoor event in northern Wisconsin. You would be thinking about climate when you plan the event in the summer rather than the winter because you have long-term knowledge that any given Saturday in the months of May to August would be a better choice for an outdoor event in Wisconsin than any given Saturday in January. However, you cannot determine the specific day that the event should be held on because it is difficult to accurately predict the weather on a specific day. Climate can be considered “average” weather.
Global Climate Change
Climate change can be understood by approaching three areas of study:
• current and past global climate change
• causes of past and present-day global climate change
• ancient and current results of climate change
It is helpful to keep these three different aspects of climate change clearly separated when consuming media reports about global climate change. It is common for reports and discussions about global climate change to confuse the data showing that Earth’s climate is changing with the factors that drive this climate change.
Evidence for Global Climate Change
Since scientists cannot go back in time to directly measure climatic variables, such as average temperature and precipitation, they must instead indirectly measure temperature. To do this, scientists rely on historical evidence of Earth’s past climate.
Antarctic ice cores are a key example of such evidence. These ice cores are samples of polar ice obtained by means of drills that reach thousands of meters into ice sheets or high mountain glaciers. Viewing the ice cores is like traveling backwards through time; the deeper the sample, the earlier the time period. Trapped within the ice are bubbles of air and other biological evidence that can reveal temperature and carbon dioxide data. Antarctic ice cores have been collected and analyzed to indirectly estimate the temperature of the Earth over the past 400,000 years (Figure \(1\)a). The 0 °C on this graph refers to the long-term average. Temperatures that are greater than 0 °C exceed Earth’s long-term average temperature. Conversely, temperatures that are less than 0 °C are less than Earth’s average temperature. This figure shows that there have been periodic cycles of increasing and decreasing temperature.
Before the late 1800s, the Earth has been as much as 9 °C cooler and about 3 °C warmer. Note that the graph in Figure \(1\)b shows that the atmospheric concentration of carbon dioxide has also risen and fallen in periodic cycles; note the relationship between carbon dioxide concentration and temperature. Figure \(1\)b shows that carbon dioxide levels in the atmosphere have historically cycled between 180 and 300 parts per million (ppm) by volume.
Figure \(1\)a does not show the last 2,000 years with enough detail to compare the changes of Earth’s temperature during the last 400,000 years with the temperature change that has occurred in the more recent past. Two significant temperature anomalies, or irregularities, have occurred in the last 2000 years. These are the Medieval Climate Anomaly (or the Medieval Warm Period) and the Little Ice Age. A third temperature anomaly aligns with the Industrial Era. The Medieval Climate Anomaly occurred between 900 and 1300 AD. During this time period, many climate scientists think that slightly warmer weather conditions prevailed in many parts of the world; the higher-than-average temperature changes varied between 0.10 °C and 0.20 °C above the norm. Although 0.10 °C does not seem large enough to produce any noticeable change, it did free seas of ice. Because of this warming, the Vikings were able to colonize Greenland.
The Little Ice Age was a cold period that occurred between 1550 AD and 1850 AD. During this time, a slight cooling of a little less than 1 °C was observed in North America, Europe, and possibly other areas of the Earth. This 1 °C change in global temperature is a seemingly small deviation in temperature (as was observed during the Medieval Climate Anomaly); however, it also resulted in noticeable changes. Historical accounts reveal a time of exceptionally harsh winters with much snow and frost.
The Industrial Revolution, which began around 1750, was characterized by changes in much of human society. Advances in agriculture increased the food supply, which improved the standard of living for people in Europe and the United States. New technologies were invented and provided jobs and cheaper goods. These new technologies were powered using fossil fuels, especially coal. The Industrial Revolution starting in the early nineteenth century ushered in the beginning of the Industrial Era. When a fossil fuel is burned, carbon dioxide is released. With the beginning of the Industrial Era, atmospheric carbon dioxide began to rise (Figure \(2\)).
Current and Past Drivers of Global Climate Change
Since it is not possible to go back in time to directly observe and measure climate, scientists use indirect evidence to determine the drivers, or factors, that may be responsible for climate change. The indirect evidence includes data collected using ice cores, boreholes (a narrow shaft bored into the ground), tree rings, glacier lengths, pollen remains, and ocean sediments. The data shows a correlation between the timing of temperature changes and drivers of climate change: before the Industrial Era (pre-1780), there were three drivers of climate change that were not related to human activity or atmospheric gases. The first of these is the Milankovitch cycles. The Milankovitch cycles describe the effects of slight changes in the Earth’s orbit on Earth’s climate. The length of the Milankovitch cycles ranges between 19,000 and 100,000 years. In other words, one could expect to see some predictable changes in the Earth’s climate associated with changes in the Earth’s orbit at a minimum of every 19,000 years.
The variation in the sun’s intensity is the second natural factor responsible for climate change. Solar intensity is the amount of solar power or energy the sun emits in a given amount of time. There is a direct relationship between solar intensity and temperature. As solar intensity increases (or decreases), the Earth’s temperature correspondingly increases (or decreases). Changes in solar intensity have been proposed as one of several possible explanations for the Little Ice Age.
Finally, volcanic eruptions are a third natural driver of climate change. Volcanic eruptions can last a few days, but the solids and gases released during an eruption can influence the climate over a period of a few years, causing short-term climate changes. The gases and solids released by volcanic eruptions can include carbon dioxide, water vapor, sulfur dioxide, hydrogen sulfide, hydrogen, and carbon monoxide. Generally, volcanic eruptions cool the climate. This occurred in 1783 when volcanos in Iceland erupted and caused the release of large volumes of sulfuric oxide. This led to haze-effect cooling, a global phenomenon that occurs when dust, ash, or other suspended particles block out sunlight and trigger lower global temperatures as a result; haze-effect cooling usually extends for one or more years. In Europe and North America, haze-effect cooling produced some of the lowest average winter temperatures on record in 1783 and 1784.
Greenhouse gases are probably the most significant drivers of the climate. When heat energy from the sun strikes the Earth, gases known as greenhouse gases trap the heat in the atmosphere, as do the glass panes of a greenhouse keep heat from escaping. The greenhouse gases that affect Earth include carbon dioxide, methane, water vapor, nitrous oxide, and ozone. Approximately half of the radiation from the sun passes through these gases in the atmosphere and strikes the Earth. This radiation is converted into thermal radiation on the Earth’s surface, and then a portion of that energy is re-radiated back into the atmosphere. Greenhouse gases, however, reflect much of the thermal energy back to the Earth’s surface. The more greenhouse gases there are in the atmosphere, the more thermal energy is reflected back to the Earth’s surface. Greenhouse gases absorb and emit radiation and are an important factor in the greenhouse effect: the warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere.
Evidence supports the relationship between atmospheric concentrations of carbon dioxide and temperature: as carbon dioxide rises, global temperature rises. Since 1950, the concentration of atmospheric carbon dioxide has increased from about 280 ppm to 382 ppm in 2006. In 2011, the atmospheric carbon dioxide concentration was 392 ppm. However, the planet would not be inhabitable by current life forms if water vapor did not produce its drastic greenhouse warming effect.
Scientists look at patterns in data and try to explain differences or deviations from these patterns. The atmospheric carbon dioxide data reveal a historical pattern of carbon dioxide increasing and decreasing, cycling between a low of 180 ppm and a high of 300 ppm. Scientists have concluded that it took around 50,000 years for the atmospheric carbon dioxide level to increase from its low minimum concentration to its higher maximum concentration. However, starting recently, atmospheric carbon dioxide concentrations have increased beyond the historical maximum of 300 ppm. The current increases in atmospheric carbon dioxide have happened very quickly—in a matter of hundreds of years rather than thousands of years. What is the reason for this difference in the rate of change and the amount of increase in carbon dioxide? A key factor that must be recognized when comparing the historical data and the current data is the presence of modern human society; no other driver of climate change has yielded changes in atmospheric carbon dioxide levels at this rate or to this magnitude.
Human activity releases carbon dioxide and methane, two of the most important greenhouse gases, into the atmosphere in several ways. The primary mechanism that releases carbon dioxide is the burning of fossil fuels, such as gasoline, coal, and natural gas (Figure \(3\)). Deforestation, cement manufacture, animal agriculture, the clearing of land, and the burning of forests are other human activities that release carbon dioxide. Methane (CH4) is produced when bacteria break down organic matter under anaerobic conditions. Anaerobic conditions can happen when organic matter is trapped underwater (such as in rice paddies) or in the intestines of herbivores. Methane can also be released from natural gas fields and the decomposition that occurs in landfills. Another source of methane is the melting of clathrates. Clathrates are frozen chunks of ice and methane found at the bottom of the ocean. When water warms, these chunks of ice melt and methane is released. As the ocean’s water temperature increases, the rate at which clathrates melt is increasing, releasing even more methane. This leads to increased levels of methane in the atmosphere, which further accelerates the rate of global warming. This is an example of the positive feedback loop that is leading to the rapid rate of increase of global temperatures.
Documented Results of Climate Change: Past and Present
Scientists have geological evidence of the consequences of long-ago climate change. Modern-day phenomena such as retreating glaciers and melting polar ice cause a continual rise in sea level. Meanwhile, changes in climate can negatively affect organisms.
Geological Climate Change
Global warming has been associated with at least one planet-wide extinction event during the geological past. The Permian extinction event occurred about 251 million years ago toward the end of the roughly 50-million-year-long geological time span known as the Permian period. This geologic time period was one of the three warmest periods in Earth’s geologic history. Scientists estimate that approximately 70 percent of the terrestrial plant and animal species and 84 percent of marine species became extinct, vanishing forever near the end of the Permian period. Organisms that had adapted to wet and warm climatic conditions, such as annual rainfall of 300–400 cm (118–157 in) and 20 °C–30 °C (68 °F–86 °F) in the tropical wet forest, may not have been able to survive the Permian climate change.
Link to Learning
Watch this NASA video to discover the mixed effects of global warming on plant growth. While scientists found that warmer temperatures in the 1980s and 1990s caused an increase in plant productivity, this advantage has since been counteracted by more frequent droughts.
Present Climate Change
A number of global events have occurred that may be attributed to climate change during our lifetimes. Glacier National Park in Montana is undergoing the retreat of many of its glaciers, a phenomenon known as glacier recession. In 1850, the area contained approximately 150 glaciers. By 2010, however, the park contained only about 24 glaciers greater than 25 acres in size. One of these glaciers is the Grinnell Glacier (Figure \(4\)) at Mount Gould. Between 1966 and 2005, the size of Grinnell Glacier shrank by 40 percent. Similarly, the mass of the ice sheets in Greenland and the Antarctic is decreasing: Greenland lost 150–250 km3 of ice per year between 2002 and 2006. In addition, the size and thickness of the Arctic sea ice is decreasing.
This loss of ice is leading to increases in the global sea level. On average, the sea is rising at a rate of 1.8 mm per year. However, between 1993 and 2010 the rate of sea level increase ranged between 2.9 and 3.4 mm per year. A variety of factors affect the volume of water in the ocean, including the temperature of the water (the density of water is related to its temperature) and the amount of water found in rivers, lakes, glaciers, polar ice caps, and sea ice. As glaciers and polar ice caps melt, there is a significant contribution of liquid water that was previously frozen.
In addition to some abiotic conditions changing in response to climate change, many organisms are also being affected by the changes in temperature. Temperature and precipitation play key roles in determining the geographic distribution and phenology of plants and animals. (Phenology is the study of the effects of climatic conditions on the timing of periodic lifecycle events, such as flowering in plants or migration in birds.) Researchers have shown that 385 plant species in Great Britain are flowering 4.5 days sooner than was recorded earlier during the previous 40 years. In addition, insect-pollinated species were more likely to flower earlier than wind-pollinated species. The impact of changes in flowering date would be mitigated if the insect pollinators emerged earlier. This mismatched timing of plants and pollinators could result in injurious ecosystem effects because, for continued survival, insect-pollinated plants must flower when their pollinators are present.
Summary
The Earth has gone through periodic cycles of increases and decreases in temperature. During the past 2000 years, the Medieval Climate Anomaly was a warmer period, while the Little Ice Age was unusually cool. Both of these irregularities can be explained by natural causes of changes in climate, and, although the temperature changes were small, they had significant effects. Natural drivers of climate change include Milankovitch cycles, changes in solar activity, and volcanic eruptions. None of these factors, however, leads to rapid increases in global temperature or sustained increases in carbon dioxide. The burning of fossil fuels is an important source of greenhouse gases, which plays a major role in the greenhouse effect. Long ago, global warming resulted in the Permian extinction: a large-scale extinction event that is documented in the fossil record. Currently, modern-day climate change is associated with the increased melting of glaciers and polar ice sheets, resulting in a gradual increase in sea level. Plants and animals can also be affected by global climate change when the timing of seasonal events, such as flowering or pollination, is affected by global warming.
Glossary
clathrates
frozen chunks of ice and methane found at the bottom of the ocean
climate
long-term, predictable atmospheric conditions present in a specific area
global climate change
altered global weather patterns, including a worldwide increase in temperature, due largely to rising levels of atmospheric carbon dioxide
greenhouse effect
warming of Earth due to carbon dioxide and other greenhouse gases in the atmosphere
greenhouse gases
atmospheric gases such as carbon dioxide and methane that absorb and emit radiation, thus trapping heat in Earth’s atmosphere
haze-effect cooling
effect of the gases and solids from a volcanic eruption on global climate
Milankovitch cycles
cyclic changes in the Earth's orbit that may affect climate
solar intensity
amount of solar power energy the sun emits in a given amount of time
weather
conditions of the atmosphere during a short period of time | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/57%3A_The_Biosphere_and_Human_Impacts/57.06%3A_Human_Impacts_on_the_Biosphere-_Climate_Change/57.6.01%3A_Climate_and_the_Effects_of_Global_Climate_Change.txt |
• 58.1: Overview of The Biodiversity Crisis
Scientists generally accept that the term biodiversity describes the number and kinds of species in a location or on the planet. Species can be difficult to define, but most biologists still feel comfortable with the concept and are able to identify and count eukaryotic species in most contexts. Biologists have also identified alternate measures of biodiversity, some of which are important for planning how to preserve biodiversity.
• 58.2: The Value of Biodiversity
Agriculture began after early hunter-gatherer societies first settled in one place and heavily modified their immediate environment. This cultural transition has made it difficult for humans to recognize their dependence on undomesticated living things on the planet. Biologists recognize the human species is embedded in ecosystems and is dependent on them, just as every other species on the planet is dependent.
• 58.3: Factors Responsible for Extinction
The core threat to biodiversity on the planet, and therefore a threat to human welfare, is the combination of human population growth and resource exploitation. The human population requires resources to survive and grow, and those resources are being removed unsustainably from the environment. The three greatest proximate threats to biodiversity are habitat loss, overharvesting, and introduction of exotic species.
• 58.4: An Evolutionary Perspective on the Biodiversity Crisis
• 58.5: Approaches for Preserving Endangered Species and Ecosystems
Preserving biodiversity is an extraordinary challenge that must be met by greater understanding of biodiversity itself, changes in human behavior and beliefs, and various preservation strategies.
58: Conservation Biology
Skills to Develop
• Define biodiversity
• Describe biodiversity as the equilibrium of naturally fluctuating rates of extinction and speciation
• Identify historical causes of high extinction rates in Earth’s history
Traditionally, ecologists have measured biodiversity, a general term for the variety present in the biosphere, by taking into account both the number of species and their commonness. Biodiversity can be estimated at a number of levels of organization of living things. These estimation indexes, which came from information theory, are most useful as a first step in quantifying biodiversity between and within ecosystems; they are less useful when the main concern among conservation biologists is simply the loss of biodiversity. However, biologists recognize that measures of biodiversity, in terms of species diversity, may help focus efforts to preserve the biologically or technologically important elements of biodiversity.
The Lake Victoria cichlids provide an example through which we can begin to understand biodiversity. The biologists studying cichlids in the 1980s discovered hundreds of cichlid species representing a variety of specializations to particular habitat types and specific feeding strategies: eating plankton floating in the water, scraping and then eating algae from rocks, eating insect larvae from the bottom, and eating the eggs of other species of cichlid. The cichlids of Lake Victoria are the product of an adaptive radiation. An adaptive radiation is a rapid (less than three million years in the case of the Lake Victoria cichlids) branching through speciation of a phylogenetic tree into many closely related species; typically, the species “radiate” into different habitats and niches. The Galápagos finches are an example of a modest adaptive radiation with 15 species. The cichlids of Lake Victoria are an example of a spectacular adaptive radiation that includes about 500 species.
At the time biologists were making this discovery, some species began to quickly disappear. A culprit in these declines was a species of large fish that was introduced to Lake Victoria by fisheries to feed the people living around the lake. The Nile perch was introduced in 1963, but lay low until the 1980s when its populations began to surge. The Nile perch population grew by consuming cichlids, driving species after species to the point of extinction (the disappearance of a species). In fact, there were several factors that played a role in the extinction of perhaps 200 cichlid species in Lake Victoria: the Nile perch, declining lake water quality due to agriculture and land clearing on the shores of Lake Victoria, and increased fishing pressure. Scientists had not even catalogued all of the species present—so many were lost that were never named. The diversity is now a shadow of what it once was.
The cichlids of Lake Victoria are a thumbnail sketch of contemporary rapid species loss that occurs all over Earth and is caused by human activity. Extinction is a natural process of macroevolution that occurs at the rate of about one out of 1 million species becoming extinct per year. The fossil record reveals that there have been five periods of mass extinction in history with much higher rates of species loss, and the rate of species loss today is comparable to those periods of mass extinction. However, there is a major difference between the previous mass extinctions and the current extinction we are experiencing: human activity. Specifically, three human activities have a major impact: destruction of habitat, introduction of exotic species, and over-harvesting. Predictions of species loss within the next century, a tiny amount of time on geological timescales, range from 10 percent to 50 percent. Extinctions on this scale have only happened five other times in the history of the planet, and they have been caused by cataclysmic events that changed the course of the history of life in each instance. Earth is now in one of those times.
Types of Biodiversity
Scientists generally accept that the term biodiversity describes the number and kinds of species in a location or on the planet. Species can be difficult to define, but most biologists still feel comfortable with the concept and are able to identify and count eukaryotic species in most contexts. Biologists have also identified alternate measures of biodiversity, some of which are important for planning how to preserve biodiversity.
Genetic diversity is one of those alternate concepts. Genetic diversity or variation is the raw material for adaptation in a species. A species’ future potential for adaptation depends on the genetic diversity held in the genomes of the individuals in populations that make up the species. The same is true for higher taxonomic categories. A genus with very different types of species will have more genetic diversity than a genus with species that look alike and have similar ecologies. If there were a choice between one of these genera of species being preserved, the one with the greatest potential for subsequent evolution is the most genetically diverse one. It would be ideal not to have to make such choices, but increasingly this may be the norm.
Many genes code for proteins, which in turn carry out the metabolic processes that keep organisms alive and reproducing. Genetic diversity can be measured as chemical diversity in that different species produce a variety of chemicals in their cells, both the proteins as well as the products and byproducts of metabolism. This chemical diversity has potential benefit for humans as a source of pharmaceuticals, so it provides one way to measure diversity that is important to human health and welfare.
Humans have generated diversity in domestic animals, plants, and fungi. This diversity is also suffering losses because of migration, market forces, and increasing globalism in agriculture, especially in heavily populated regions such as China, India, and Japan. The human population directly depends on this diversity as a stable food source, and its decline is troubling biologists and agricultural scientists.
It is also useful to define ecosystem diversity, meaning the number of different ecosystems on the planet or in a given geographic area (Figure \(1\)). Whole ecosystems can disappear even if some of the species might survive by adapting to other ecosystems. The loss of an ecosystem means the loss of interactions between species, the loss of unique features of coadaptation, and the loss of biological productivity that an ecosystem is able to create. An example of a largely extinct ecosystem in North America is the prairie ecosystem. Prairies once spanned central North America from the boreal forest in northern Canada down into Mexico. They are now all but gone, replaced by crop fields, pasture lands, and suburban sprawl. Many of the species survive, but the hugely productive ecosystem that was responsible for creating the most productive agricultural soils is now gone. As a consequence, soils are disappearing or must be maintained at greater expense.
Current Species Diversity
Despite considerable effort, knowledge of the species that inhabit the planet is limited. A recent estimate suggests that the eukaryote species for which science has names, about 1.5 million species, account for less than 20 percent of the total number of eukaryote species present on the planet (8.7 million species, by one estimate). Estimates of numbers of prokaryotic species are largely guesses, but biologists agree that science has only begun to catalog their diversity. Even with what is known, there is no central repository of names or samples of the described species; therefore, there is no way to be sure that the 1.5 million descriptions is an accurate number. It is a best guess based on the opinions of experts in different taxonomic groups. Given that Earth is losing species at an accelerating pace, science is very much in the place it was with the Lake Victoria cichlids: knowing little about what is being lost. Table \(1\) presents recent estimates of biodiversity in different groups.
Table \(1\): Estimates of the described and predicted species by Taxonomic group
Mora et al. 20111 Chapman 20092 Groombridge & Jenkins 20023
Described Predicted Described Predicted Described Predicted
Animalia 1,124,516 9,920,000 1,424,153 6,836,330 1,225,500 10,820,000
Chromista 17,892 34,900 25,044 200,500
Fungi 44,368 616,320 98,998 1,500,000 72,000 1,500,000
Plantae 224,244 314,600 310,129 390,800 270,000 320,000
Protozoa 16,236 72,800 28,871 1,000,000 80,000 600,000
Prokaryotes 10,307 1,000,000 10,175
Total 1,438,769 10,960,000 1,897,502 10,897,630 1,657,675 13,240,000
There are various initiatives to catalog described species in accessible ways, and the internet is facilitating that effort. Nevertheless, it has been pointed out that at the current rate of species description, which according to the State of Observed Species Report is 17,000 to 20,000 new species per year, it will take close to 500 years to finish describing life on this planet.4 Over time, the task becomes both increasingly impossible and increasingly easier as extinction removes species from the planet.
Naming and counting species may seem an unimportant pursuit given the other needs of humanity, but it is not simply an accounting. Describing species is a complex process by which biologists determine an organism’s unique characteristics and whether or not that organism belongs to any other described species. It allows biologists to find and recognize the species after the initial discovery, and allows them to follow up on questions about its biology. In addition, the unique characteristics of each species make it potentially valuable to humans or other species on which humans depend. Understanding these characteristics is the value of finding and naming species.
Patterns of Biodiversity
Biodiversity is not evenly distributed on Earth. Lake Victoria contained almost 500 species of cichlids alone, ignoring the other fish families present in the lake. All of these species were found only in Lake Victoria; therefore, the 500 species of cichlids were endemic. Endemic species are found in only one location. Endemics with highly restricted distributions are particularly vulnerable to extinction. Higher taxonomic levels, such as genera and families, can also be endemic. Lake Huron contains about 79 species of fish, all of which are found in many other lakes in North America. What accounts for the difference in fish diversity in these two lakes? Lake Victoria is a tropical lake, while Lake Huron is a temperate lake. Lake Huron in its present form is only about 7,000 years old, while Lake Victoria in its present form is about 15,000 years old. Biogeographers have suggested these two factors, latitude and age, are two of several hypotheses to explain biodiversity patterns on the planet.
Career Connection: Biogeographer
Biogeography is the study of the distribution of the world’s species—both in the past and in the present. The work of biogeographers is critical to understanding our physical environment, how the environment affects species, and how environmental changes impact the distribution of a species; it has also been critical to developing evolutionary theory. Biogeographers need to understand both biology and ecology. They also need to be well-versed in evolutionary studies, soil science, and climatology.
There are three main fields of study under the heading of biogeography: ecological biogeography, historical biogeography (called paleobiogeography), and conservation biogeography. Ecological biogeography studies the current factors affecting the distribution of plants and animals. Historical biogeography, as the name implies, studies the past distribution of species. Conservation biogeography, on the other hand, is focused on the protection and restoration of species based upon known historical and current ecological information. Each of these fields considers both zoogeography and phytogeography—the past and present distribution of animals and plants.
One of the oldest observed patterns in ecology is that species biodiversity in almost every taxonomic group increases as latitude declines. In other words, biodiversity increases closer to the equator (Figure \(2\)).
It is not yet clear why biodiversity increases closer to the equator, but hypotheses include the greater age of the ecosystems in the tropics versus temperate regions that were largely devoid of life or drastically impoverished during the last glaciation. The idea is that greater age provides more time for speciation. Another possible explanation is the increased energy the tropics receive from the sun versus the decreased energy that temperate and polar regions receive. It is not entirely clear how greater energy input could translate into more species. The complexity of tropical ecosystems may promote speciation by increasing the heterogeneity, or number of ecological niches, in the tropics relative to higher latitudes. The greater heterogeneity provides more opportunities for coevolution, specialization, and perhaps greater selection pressures leading to population differentiation. However, this hypothesis suffers from some circularity—ecosystems with more species encourage speciation, but how did they get more species to begin with? The tropics have been perceived as being more stable than temperate regions, which have a pronounced climate and day-length seasonality. The tropics have their own forms of seasonality, such as rainfall, but they are generally assumed to be more stable environments and this stability might promote speciation.
Regardless of the mechanisms, it is certainly true that all levels of biodiversity are greatest in the tropics. Additionally, the rate of endemism is highest, and there are more biodiversity hotspots. However, this richness of diversity also means that knowledge of species is lowest, and there is a high potential for biodiversity loss.
Conservation of Biodiversity
In 1988, British environmentalist Norman Myers developed a conservation concept to identify areas rich in species and at significant risk for species loss: biodiversity hotspots. Biodiversity hotspots are geographical areas that contain high numbers of endemic species. The purpose of the concept was to identify important locations on the planet for conservation efforts, a kind of conservation triage. By protecting hotspots, governments are able to protect a larger number of species. The original criteria for a hotspot included the presence of 1500 or more endemic plant species and 70 percent of the area disturbed by human activity. There are now 34 biodiversity hotspots (Figure \(3\)) containing large numbers of endemic species, which include half of Earth’s endemic plants.
Biodiversity Change through Geological Time
The number of species on the planet, or in any geographical area, is the result of an equilibrium of two evolutionary processes that are ongoing: speciation and extinction. Both are natural “birth” and “death” processes of macroevolution. When speciation rates begin to outstrip extinction rates, the number of species will increase; likewise, the number of species will decrease when extinction rates begin to overtake speciation rates. Throughout Earth’s history, these two processes have fluctuated—sometimes leading to dramatic changes in the number of species on Earth as reflected in the fossil record (Figure \(4\)).
Paleontologists have identified five strata in the fossil record that appear to show sudden and dramatic (greater than half of all extant species disappearing from the fossil record) losses in biodiversity. These are called mass extinctions. There are many lesser, yet still dramatic, extinction events, but the five mass extinctions have attracted the most research. An argument can be made that the five mass extinctions are only the five most extreme events in a continuous series of large extinction events throughout the Phanerozoic (since 542 million years ago). In most cases, the hypothesized causes are still controversial; however, the most recent event seems clear.
The Five Mass Extinctions
The fossil record of the mass extinctions was the basis for defining periods of geological history, so they typically occur at the transition point between geological periods. The transition in fossils from one period to another reflects the dramatic loss of species and the gradual origin of new species. These transitions can be seen in the rock strata. Table \(2\) provides data on the five mass extinctions.
This table shows the names and dates for the five mass extinctions in Earth’s history.
Table \(2\): Mass extinctions
Geological Period Mass Extinction Name Time (millions of years ago)
Ordovician–Silurian end-Ordovician O–S 450–440
Late Devonian end-Devonian 375–360
Permian–Triassic end-Permian 251
Triassic–Jurassic end-Triassic 205
Cretaceous–Paleogene end-Cretaceous K–Pg (K–T) 65.5
The Ordovician-Silurian extinction event is the first recorded mass extinction and the second largest. During this period, about 85 percent of marine species (few species lived outside the oceans) became extinct. The main hypothesis for its cause is a period of glaciation and then warming. The extinction event actually consists of two extinction events separated by about 1 million years. The first event was caused by cooling, and the second event was due to the subsequent warming. The climate changes affected temperatures and sea levels. Some researchers have suggested that a gamma-ray burst, caused by a nearby supernova, is a possible cause of the Ordovician-Silurian extinction. The gamma-ray burst would have stripped away the Earth’s ozone layer causing intense ultraviolet radiation from the sun and may account for climate changes observed at the time. The hypothesis is speculative, but extraterrestrial influences on Earth’s history are an active line of research. Recovery of biodiversity after the mass extinction took from 5 to 20 million years, depending on the location.
The late Devonian extinction may have occurred over a relatively long period of time. It appears to have affected marine species and not the plants or animals inhabiting terrestrial habitats. The causes of this extinction are poorly understood.
The end-Permian extinction was the largest in the history of life. Indeed, an argument could be made that Earth nearly became devoid of life during this extinction event. The planet looked very different before and after this event. Estimates are that 96 percent of all marine species and 70 percent of all terrestrial species were lost. It was at this time, for example, that the trilobites, a group that survived the Ordovician–Silurian extinction, became extinct. The causes for this mass extinction are not clear, but the leading suspect is extended and widespread volcanic activity that led to a runaway global-warming event. The oceans became largely anoxic, suffocating marine life. Terrestrial tetrapod diversity took 30 million years to recover after the end-Permian extinction. The Permian extinction dramatically altered Earth’s biodiversity makeup and the course of evolution.
The causes of the Triassic–Jurassic extinction event are not clear and hypotheses of climate change, asteroid impact, and volcanic eruptions have been argued. The extinction event occurred just before the breakup of the supercontinent Pangaea, although recent scholarship suggests that the extinctions may have occurred more gradually throughout the Triassic.
The causes of the end-Cretaceous extinction event are the ones that are best understood. It was during this extinction event about 65 million years ago that the dinosaurs, the dominant vertebrate group for millions of years, disappeared from the planet (with the exception of a theropod clade that gave rise to birds). Indeed, every land animal that weighed more then 25 kg became extinct. The cause of this extinction is now understood to be the result of a cataclysmic impact of a large meteorite, or asteroid, off the coast of what is now the Yucatán Peninsula. This hypothesis, proposed first in 1980, was a radical explanation based on a sharp spike in the levels of iridium (which rains down from space in meteors at a fairly constant rate but is otherwise absent on Earth’s surface) at the rock stratum that marks the boundary between the Cretaceous and Paleogene periods (Figure \(5\)). This boundary marked the disappearance of the dinosaurs in fossils as well as many other taxa. The researchers who discovered the iridium spike interpreted it as a rapid influx of iridium from space to the atmosphere (in the form of a large asteroid) rather than a slowing in the deposition of sediments during that period. It was a radical explanation, but the report of an appropriately aged and sized impact crater in 1991 made the hypothesis more believable. Now an abundance of geological evidence supports the theory. Recovery times for biodiversity after the end-Cretaceous extinction are shorter, in geological time, than for the end-Permian extinction, on the order of 10 million years.
Art Connection
Scientists measured the relative abundance of fern spores above and below the K–Pg boundary in this rock sample. Which of the following statements most likely represents their findings?
1. An abundance of fern spores from several species was found below the K–Pg boundary, but none was found above.
2. An abundance of fern spores from several species was found above the K–Pg boundary, but none was found below.
3. An abundance of fern spores was found both above and below the K–Pg boundary, but only one species was found below the boundary, and many species were found above the boundary.
4. Many species of fern spores were found both above and below the boundary, but the total number of spores was greater below the boundary.
Link to Learning
Explore this interactive website about mass extinctions.
The Pleistocene Extinction
The Pleistocene Extinction is one of the lesser extinctions, and a recent one. It is well known that the North American, and to some degree Eurasian, megafauna, or large animals, disappeared toward the end of the last glaciation period. The extinction appears to have happened in a relatively restricted time period of 10,000–12,000 years ago. In North America, the losses were quite dramatic and included the woolly mammoths (last dated about 4,000 years ago in an isolated population), mastodon, giant beavers, giant ground sloths, saber-toothed cats, and the North American camel, just to name a few. The possibility that the rapid extinction of these large animals was caused by over-hunting was first suggested in the 1900s. Research into this hypothesis continues today. It seems likely that over-hunting caused many pre-written history extinctions in many regions of the world.
In general, the timing of the Pleistocene extinctions correlated with the arrival of humans and not with climate-change events, which is the main competing hypothesis for these extinctions. The extinctions began in Australia about 40,000 to 50,000 years ago, just after the arrival of humans in the area: a marsupial lion, a giant one-ton wombat, and several giant kangaroo species disappeared. In North America, the extinctions of almost all of the large mammals occurred 10,000–12,000 years ago. All that are left are the smaller mammals such as bears, elk, moose, and cougars. Finally, on many remote oceanic islands, the extinctions of many species occurred coincident with human arrivals. Not all of the islands had large animals, but when there were large animals, they were lost. Madagascar was colonized about 2,000 years ago and the large mammals that lived there became extinct. Eurasia and Africa do not show this pattern, but they also did not experience a recent arrival of humans. Humans arrived in Eurasia hundreds of thousands of years ago after the origin of the species in Africa. This topic remains an area of active research and hypothesizing. It seems clear that even if climate played a role, in most cases human hunting precipitated the extinctions.
Present-Time Extinctions
The sixth, or Holocene, mass extinction appears to have begun earlier than previously believed and has mostly to do with the activities of Homo sapiens. Since the beginning of the Holocene period, there are numerous recent extinctions of individual species that are recorded in human writings. Most of these are coincident with the expansion of the European colonies since the 1500s.
One of the earlier and popularly known examples is the dodo bird. The dodo bird lived in the forests of Mauritius, an island in the Indian Ocean. The dodo bird became extinct around 1662. It was hunted for its meat by sailors and was easy prey because the dodo, which did not evolve with humans, would approach people without fear. Introduced pigs, rats, and dogs brought to the island by European ships also killed dodo young and eggs.
Steller's sea cow became extinct in 1768; it was related to the manatee and probably once lived along the northwest coast of North America. Steller's sea cow was first discovered by Europeans in 1741 and was hunted for meat and oil. The last sea cow was killed in 1768. That amounts to 27 years between the sea cow’s first contact with Europeans and extinction of the species.
In 1914, the last living passenger pigeon died in a zoo in Cincinnati, Ohio. This species had once darkened the skies of North America during its migrations, but it was hunted and suffered from habitat loss through the clearing of forests for farmland. In 1918, the last living Carolina parakeet died in captivity. This species was once common in the eastern United States, but it suffered from habitat loss. The species was also hunted because it ate orchard fruit when its native foods were destroyed to make way for farmland. The Japanese sea lion, which inhabited a broad area around Japan and the coast of Korea, became extinct in the 1950s due to fishermen. The Caribbean monk seal was distributed throughout the Caribbean Sea but was driven to extinction via hunting by 1952.
These are only a few of the recorded extinctions in the past 500 years. The International Union for Conservation of Nature (IUCN) keeps a list of extinct and endangered species called the Red List. The list is not complete, but it describes 380 extinct species of vertebrates after 1500 AD, 86 of which were driven extinct by overhunting or overfishing.
Estimates of Present-Time Extinction Rates
Estimates of extinction rates are hampered by the fact that most extinctions are probably happening without observation. The extinction of a bird or mammal is likely to be noticed by humans, especially if it has been hunted or used in some other way. But there are many organisms that are of less interest to humans (not necessarily of less value) and many that are undescribed.
The background extinction rate is estimated to be about one per million species per year (E/MSY). For example, assuming there are about ten million species in existence, the expectation is that ten species would become extinct each year (each year represents ten million species per year).
One contemporary extinction rate estimate uses the extinctions in the written record since the year 1500. For birds alone this method yields an estimate of 26 E/MSY. However, this value may be underestimated for three reasons. First, many species would not have been described until much later in the time period, so their loss would have gone unnoticed. Second, the number of recently extinct species is increasing because extinct species now are being described from skeletal remains. And third, some species are probably already extinct even though conservationists are reluctant to name them as such. Taking these factors into account raises the estimated extinction rate closer to 100 E/MSY. The predicted rate by the end of the century is 1500 E/MSY.
A second approach to estimating present-time extinction rates is to correlate species loss with habitat loss by measuring forest-area loss and understanding species-area relationships. The species-area relationship is the rate at which new species are seen when the area surveyed is increased. Studies have shown that the number of species present increases as the size of the island increases. This phenomenon has also been shown to hold true in other habitats as well. Turning this relationship around, if the habitat area is reduced, the number of species living there will also decline. Estimates of extinction rates based on habitat loss and species-area relationships have suggested that with about 90 percent habitat loss an expected 50 percent of species would become extinct. Species-area estimates have led to species extinction rate calculations of about 1000 E/MSY and higher. In general, actual observations do not show this amount of loss and suggestions have been made that there is a delay in extinction. Recent work has also called into question the applicability of the species-area relationship when estimating the loss of species. This work argues that the species-area relationship leads to an overestimate of extinction rates. A better relationship to use may be the endemics-area relationship. Using this method would bring estimates down to around 500 E/MSY in the coming century. Note that this value is still 500 times the background rate.
Link to Learning
Check out this interactive exploration of endangered and extinct species, their ecosystems, and the causes of the endangerment or extinction.
Summary
Biodiversity exists at multiple levels of organization and is measured in different ways depending on the goals of those taking the measurements. These measurements include numbers of species, genetic diversity, chemical diversity, and ecosystem diversity. The number of described species is estimated to be 1.5 million with about 17,000 new species being described each year. Estimates for the total number of species on Earth vary but are on the order of 10 million. Biodiversity is negatively correlated with latitude for most taxa, meaning that biodiversity is higher in the tropics. The mechanism for this pattern is not known with certainty, but several plausible hypotheses have been advanced.
Five mass extinctions with losses of more than 50 percent of extant species are observable in the fossil record. Biodiversity recovery times after mass extinctions vary, but have been up to 30 million years. Recent extinctions are recorded in written history and are the basis for one method of estimating contemporary extinction rates. The other method uses measures of habitat loss and species-area relationships. Estimates of contemporary extinction rates vary, but some rates are as high as 500 times the background rate, as determined from the fossil record, and are predicted to rise.
Art Connections
Figure \(5\): Scientists measured the relative abundance of fern spores above and below the K-Pg boundary in this rock sample. Which of the following statements most likely represents their findings?
1. An abundance of fern spores from several species was found below the K-Pg boundary, but none was found above.
2. An abundance of fern spores from several species was found above the K-Pg boundary, but none was found below.
3. An abundance of fern spores was found both above and below the K-Pg boundary, but only one species was found below the boundary , and many species were found above the boundary.
4. Many species of fern spores were found both above and below the boundary, but the total number of spores was greater below the boundary.
Answer
A. An abundance of fern spores from several species was found below the K-Pg boundary, but none was found above.
Footnotes
1. 1 Mora Camilo et al., “How Many Species Are There on Earth and in the Ocean?” PLoS Biology (2011), doi:10.1371/journal.pbio.1001127.
2. 2 Arthur D. Chapman, Numbers of Living Species in Australia and the World, 2nd ed. (Canberra, AU: Australian Biological Resources Study, 2009). www.environment.gov.au/biodiv...d-complete.pdf.
3. 3 Brian Groombridge and Martin D. Jenkins. World Atlas of Biodiversity: Earth’s Living Resources in the 21st Century. Berkeley: University of California Press, 2002.
4. 4 International Institute for Species Exploration (IISE), 2011 State of Observed Species (SOS). Tempe, AZ: IISE, 2011. Accessed May, 20, 2012. species.asu.edu/SOS.
Glossary
adaptive radiation
rapid branching through speciation of a phylogenetic tree into many closely related species
biodiversity
variety of a biological system, typically conceived as the number of species, but also applying to genes, biochemistry, and ecosystems
biodiversity hotspot
concept originated by Norman Myers to describe a geographical region with a large number of endemic species and a large percentage of degraded habitat
chemical diversity
variety of metabolic compounds in an ecosystem
ecosystem diversity
variety of ecosystems
endemic species
species native to one place
extinction
disappearance of a species from Earth; local extinction is the disappearance of a species from a region
extinction rate
number of species becoming extinct over time, sometimes defined as extinctions per million species–years to make numbers manageable (E/MSY)
genetic diversity
variety of genes in a species or other taxonomic group or ecosystem, the term can refer to allelic diversity or genome-wide diversity
heterogeneity
number of ecological niches
megafauna
large animals
species-area relationship
relationship between area surveyed and number of species encountered; typically measured by incrementally increasing the area of a survey and determining the cumulative numbers of species | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/58%3A_Conservation_Biology/58.01%3A_Overview_of_The_Biodiversity_Crisis.txt |
Skills to Develop
• Identify chemical diversity benefits to humans
• Identify biodiversity components that support human agriculture
• Describe ecosystem services
It may not be clear why biologists are concerned about biodiversity loss. When biodiversity loss is thought of as the extinction of the passenger pigeon, the dodo bird, and even the woolly mammoth, the loss may appear to be an emotional one. But is the loss practically important for the welfare of the human species? From the perspective of evolution and ecology, the loss of a particular individual species is unimportant (however, the loss of a keystone species can lead to ecological disaster). Extinction is a normal part of macroevolution. But the accelerated extinction rate means the loss of tens of thousands of species within our lifetimes, and it is likely to have dramatic effects on human welfare through the collapse of ecosystems and in added costs to maintain food production, clean air and water, and human health.
Agriculture began after early hunter-gatherer societies first settled in one place and heavily modified their immediate environment. This cultural transition has made it difficult for humans to recognize their dependence on undomesticated living things on the planet. Biologists recognize the human species is embedded in ecosystems and is dependent on them, just as every other species on the planet is dependent. Technology smoothes out the extremes of existence, but ultimately the human species cannot exist without its ecosystem.
Human Health
Contemporary societies that live close to the land often have a broad knowledge of the medicinal uses of plants growing in their area. Most plants produce secondary plant compounds, which are toxins used to protect the plant from insects and other animals that eat them, but some of which also work as medication. For centuries in Europe, older knowledge about the medical uses of plants was compiled in herbals—books that identified plants and their uses. Humans are not the only species to use plants for medicinal reasons: the great apes, orangutans, chimpanzees, bonobos, and gorillas have all been observed self-medicating with plants.
Modern pharmaceutical science also recognizes the importance of these plant compounds. Examples of significant medicines derived from plant compounds include aspirin, codeine, digoxin, atropine, and vincristine (Figure \(1\)). Many medicines were once derived from plant extracts but are now synthesized. It is estimated that, at one time, 25 percent of modern drugs contained at least one plant extract. That number has probably decreased to about 10 percent as natural plant ingredients are replaced by synthetic versions. Antibiotics, which are responsible for extraordinary improvements in health and lifespans in developed countries, are compounds largely derived from fungi and bacteria.
In recent years, animal venoms and poisons have excited intense research for their medicinal potential. By 2007, the FDA had approved five drugs based on animal toxins to treat diseases such as hypertension, chronic pain, and diabetes. Another five drugs are undergoing clinical trials, and at least six drugs are being used in other countries. Other toxins under investigation come from mammals, snakes, lizards, various amphibians, fish, snails, octopuses, and scorpions.
Aside from representing billions of dollars in profits, these medicines improve people’s lives. Pharmaceutical companies are actively looking for new compounds synthesized by living organisms that can function as medicine. It is estimated that 1/3 of pharmaceutical research and development is spent on natural compounds and that about 35 percent of new drugs brought to market between 1981 and 2002 were from natural compounds. The opportunities for new medications will be reduced in direct proportion to the disappearance of species.
Agricultural Diversity
Since the beginning of human agriculture more than 10,000 years ago, human groups have been breeding and selecting crop varieties. This crop diversity matched the cultural diversity of highly subdivided populations of humans. For example, potatoes were domesticated beginning around 7,000 years ago in the central Andes of Peru and Bolivia. The potatoes grown in that region belong to seven species and the number of varieties likely is in the thousands. Each variety has been bred to thrive at particular elevations and soil and climate conditions. The diversity is driven by the diverse demands of the topography, the limited movement of people, and the demands created by crop rotation for different varieties that will do well in different fields.
Potatoes are only one example of human-generated diversity. Every plant, animal, and fungus that has been cultivated by humans has been bred from original wild ancestor species into diverse varieties arising from the demands for food value, adaptation to growing conditions, and resistance to pests. The potato demonstrates a well-known example of the risks of low crop diversity: the tragic Irish potato famine when the single variety grown in Ireland became susceptible to a potato blight, wiping out the crop. The loss of the crop led to famine, death, and mass emigration. Resistance to disease is a chief benefit to maintaining crop biodiversity, and lack of diversity in contemporary crop species carries similar risks. Seed companies, which are the source of most crop varieties in developed countries, must continually breed new varieties to keep up with evolving pest organisms. These same seed companies, however, have participated in the decline of the number of varieties available as they focus on selling fewer varieties in more areas of the world.
The ability to create new crop varieties relies on the diversity of varieties available and the accessibility of wild forms related to the crop plant. These wild forms are often the source of new gene variants that can be bred with existing varieties to create varieties with new attributes. Loss of wild species related to a crop will mean the loss of potential in crop improvement. Maintaining the genetic diversity of wild species related to domesticated species ensures our continued food supply.
Since the 1920s, government agriculture departments have maintained seed banks of crop varieties as a way to maintain crop diversity. This system has flaws because, over time, seed banks are lost through accidents, and there is no way to replace them. In 2008, the Svalbard Global Seed Vault (Figure \(2\)) began storing seeds from around the world as a backup system to the regional seed banks. If a regional seed bank stores varieties in Svalbard, losses can be replaced from Svalbard. The seed vault is located deep into the rock of an arctic island. Conditions within the vault are maintained at ideal temperature and humidity for seed survival, but the deep underground location of the vault in the arctic means that failure of the vault’s systems will not compromise the climatic conditions inside the vault.
Art Connection
The Svalbard Global Seed Vault is located on Spitsbergen island in Norway, which has an arctic climate. Why might an arctic climate be good for seed storage?
Crop success is largely dependent on the quality of the soil. Although some agricultural soils are rendered sterile using controversial cultivation and chemical treatments, most contain a huge diversity of organisms that maintain nutrient cycles—breaking down organic matter into nutrient compounds that crops need for growth. These organisms also maintain soil texture that affects water and oxygen dynamics in the soil that are necessary for plant growth. If farmers had to maintain arable soil using alternate means, the cost of food would be much higher than it is now. These kinds of processes are called ecosystem services. They occur within ecosystems, such as soil ecosystems, as a result of the diverse metabolic activities of the organisms living there, but they provide benefits to human food production, drinking water availability, and breathable air.
Other key ecosystem services related to food production are plant pollination and crop pest control. Over 150 crops in the United States require pollination to produce. One estimate of the benefit of honeybee pollination within the United States is \$1.6 billion per year; other pollinators contribute up to \$6.7 billion more.
Many honeybee populations are managed by apiarists who rent out their hives’ services to farmers. Honeybee populations in North America have been suffering large losses caused by a syndrome known as colony collapse disorder, whose cause is unclear. Other pollinators include a diverse array of other bee species and various insects and birds. Loss of these species would make growing crops requiring pollination impossible, increasing dependence on other crops.
Finally, humans compete for their food with crop pests, most of which are insects. Pesticides control these competitors; however, pesticides are costly and lose their effectiveness over time as pest populations adapt. They also lead to collateral damage by killing non-pest species and risking the health of consumers and agricultural workers. Ecologists believe that the bulk of the work in removing pests is actually done by predators and parasites of those pests, but the impact has not been well studied. A review found that in 74 percent of studies that looked for an effect of landscape complexity on natural enemies of pests, the greater the complexity, the greater the effect of pest-suppressing organisms. An experimental study found that introducing multiple enemies of pea aphids (an important alfalfa pest) increased the yield of alfalfa significantly. This study shows the importance of landscape diversity via the question of whether a diversity of pests is more effective at control than one single pest; the results showed this to be the case. Loss of diversity in pest enemies will inevitably make it more difficult and costly to grow food.
Wild Food Sources
In addition to growing crops and raising animals for food, humans obtain food resources from wild populations, primarily fish populations. For approximately 1 billion people, aquatic resources provide the main source of animal protein. But since 1990, global fish production has declined. Despite considerable effort, few fisheries on the planet are managed for sustainability.
Fishery extinctions rarely lead to complete extinction of the harvested species, but rather to a radical restructuring of the marine ecosystem in which a dominant species is so over-harvested that it becomes a minor player, ecologically. In addition to humans losing the food source, these alterations affect many other species in ways that are difficult or impossible to predict. The collapse of fisheries has dramatic and long-lasting effects on local populations that work in the fishery. In addition, the loss of an inexpensive protein source to populations that cannot afford to replace it will increase the cost of living and limit societies in other ways. In general, the fish taken from fisheries have shifted to smaller species as larger species are fished to extinction. The ultimate outcome could clearly be the loss of aquatic systems as food sources.
Psychological and Moral Value
Finally, it has been argued that humans benefit psychologically from living in a biodiverse world. A chief proponent of this idea is entomologist E. O. Wilson. He argues that human evolutionary history has adapted us to live in a natural environment and that built environments generate stressors that affect human health and well-being. There is considerable research into the psychological regenerative benefits of natural landscapes that suggests the hypothesis may hold some truth. In addition, there is a moral argument that humans have a responsibility to inflict as little harm as possible on other species.
Summary
Humans use many compounds that were first discovered or derived from living organisms as medicines: secondary plant compounds, animal toxins, and antibiotics produced by bacteria and fungi. More medicines are expected to be discovered in nature. Loss of biodiversity will impact the number of pharmaceuticals available to humans.
Crop diversity is a requirement for food security, and it is being lost. The loss of wild relatives to crops also threatens breeders’ abilities to create new varieties. Ecosystems provide ecosystem services that support human agriculture: pollination, nutrient cycling, pest control, and soil development and maintenance. Loss of biodiversity threatens these ecosystem services and risks making food production more expensive or impossible. Wild food sources are mainly aquatic, but few are being managed for sustainability. Fisheries’ ability to provide protein to human populations is threatened when extinction occurs.
Biodiversity may provide important psychological benefits to humans. Additionally, there are moral arguments for the maintenance of biodiversity.
Art Connections
Figure \(2\): The Svalbard Global Seed Vault is located on Spitsbergen island in Norway, which has an arctic climate. Why might an arctic climate be good for seed storage?
Answer
The ground is permanently frozen so the seeds will keep even if the electricity fails.
Glossary
secondary plant compound
compound produced as byproducts of plant metabolic processes that is usually toxic, but is sequestered by the plant to defend against herbivores | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/58%3A_Conservation_Biology/58.02%3A_The_Value_of_Biodiversity.txt |
Skills to Develop
• Identify significant threats to biodiversity
• Explain the effects of habitat loss, exotic species, and hunting on biodiversity
• Identify the early and predicted effects of climate change on biodiversity
The core threat to biodiversity on the planet, and therefore a threat to human welfare, is the combination of human population growth and resource exploitation. The human population requires resources to survive and grow, and those resources are being removed unsustainably from the environment. The three greatest proximate threats to biodiversity are habitat loss, overharvesting, and introduction of exotic species. The first two of these are a direct result of human population growth and resource use. The third results from increased mobility and trade. A fourth major cause of extinction, anthropogenic climate change, has not yet had a large impact, but it is predicted to become significant during this century. Global climate change is also a consequence of human population needs for energy and the use of fossil fuels to meet those needs (Figure \(1\)). Environmental issues, such as toxic pollution, have specific targeted effects on species, but they are not generally seen as threats at the magnitude of the others.
Habitat Loss
Humans rely on technology to modify their environment and replace certain functions that were once performed by the natural ecosystem. Other species cannot do this. Elimination of their ecosystem—whether it is a forest, a desert, a grassland, a freshwater estuarine, or a marine environment—will kill the individuals in the species. Remove the entire habitat within the range of a species and, unless they are one of the few species that do well in human-built environments, the species will become extinct. Human destruction of habitats accelerated in the latter half of the twentieth century. Consider the exceptional biodiversity of Sumatra: it is home to one species of orangutan, a species of critically endangered elephant, and the Sumatran tiger, but half of Sumatra’s forest is now gone. The neighboring island of Borneo, home to the other species of orangutan, has lost a similar area of forest. Forest loss continues in protected areas of Borneo. The orangutan in Borneo is listed as endangered by the International Union for Conservation of Nature (IUCN), but it is simply the most visible of thousands of species that will not survive the disappearance of the forests of Borneo. The forests are removed for timber and to plant palm oil plantations (Figure \(2\)). Palm oil is used in many products including food products, cosmetics, and biodiesel in Europe. A five-year estimate of global forest cover loss for the years 2000–2005 was 3.1 percent. In the humid tropics where forest loss is primarily from timber extraction, 272,000 km2 was lost out of a global total of 11,564,000 km2 (or 2.4 percent). In the tropics, these losses certainly also represent the extinction of species because of high levels of endemism.
Everyday Connection: Preventing Habitat Destruction with Wise Wood Choices
Most consumers do not imagine that the home improvement products they buy might be contributing to habitat loss and species extinctions. Yet the market for illegally harvested tropical timber is huge, and the wood products often find themselves in building supply stores in the United States. One estimate is that 10 percent of the imported timber stream in the United States, which is the world’s largest consumer of wood products, is potentially illegally logged. In 2006, this amounted to \$3.6 billion in wood products. Most of the illegal products are imported from countries that act as intermediaries and are not the originators of the wood.
How is it possible to determine if a wood product, such as flooring, was harvested sustainably or even legally? The Forest Stewardship Council (FSC) certifies sustainably harvested forest products, therefore, looking for their certification on flooring and other hardwood products is one way to ensure that the wood has not been taken illegally from a tropical forest. Certification applies to specific products, not to a producer; some producers’ products may not have certification while other products are certified. While there are other industry-backed certifications other than the FSC, these are unreliable due to lack of independence from the industry. Another approach is to buy domestic wood species. While it would be great if there was a list of legal versus illegal wood products, it is not that simple. Logging and forest management laws vary from country to country; what is illegal in one country may be legal in another. Where and how a product is harvested and whether the forest from which it comes is being maintained sustainably all factor into whether a wood product will be certified by the FSC. It is always a good idea to ask questions about where a wood product came from and how the supplier knows that it was harvested legally.
Habitat destruction can affect ecosystems other than forests. Rivers and streams are important ecosystems and are frequently modified through land development and from damming or water removal. Damming of rivers affects the water flow and access to all parts of a river. Differing flow regimes can reduce or eliminate populations that are adapted to these changes in flow patterns. For example, an estimated 91percent of river lengths in the United States have been developed: they have modifications like dams, to create energy or store water; levees, to prevent flooding; or dredging or rerouting, to create land that is more suitable for human development. Many fish species in the United States, especially rare species or species with restricted distributions, have seen declines caused by river damming and habitat loss. Research has confirmed that species of amphibians that must carry out parts of their life cycles in both aquatic and terrestrial habitats have a greater chance of suffering population declines and extinction because of the increased likelihood that one of their habitats or access between them will be lost.
Overharvesting
Overharvesting is a serious threat to many species, but particularly to aquatic species. There are many examples of regulated commercial fisheries monitored by fisheries scientists that have nevertheless collapsed. The western Atlantic cod fishery is the most spectacular recent collapse. While it was a hugely productive fishery for 400 years, the introduction of modern factory trawlers in the 1980s and the pressure on the fishery led to it becoming unsustainable. The causes of fishery collapse are both economic and political in nature. Most fisheries are managed as a common (shared) resource even when the fishing territory lies within a country’s territorial waters. Common resources are subject to an economic pressure known as the tragedy of the commons in which essentially no fisher has a motivation to exercise restraint in harvesting a fishery when it is not owned by that fisher. The natural outcome of harvests of resources held in common is their overexploitation. While large fisheries are regulated to attempt to avoid this pressure, it still exists in the background. This overexploitation is exacerbated when access to the fishery is open and unregulated and when technology gives fishers the ability to overfish. In a few fisheries, the biological growth of the resource is less than the potential growth of the profits made from fishing if that time and money were invested elsewhere. In these cases—whales are an example—economic forces will always drive toward fishing the population to extinction.
Link to Learning
Explore a U.S. Fish & Wildlife Service interactive map of critical habitat for endangered and threatened species in the United States. To begin, select “Visit the online mapper.”
For the most part, fishery extinction is not equivalent to biological extinction—the last fish of a species is rarely fished out of the ocean. At the same time, fishery extinction is still harmful to fish species and their ecosystems. There are some instances in which true extinction is a possibility. Whales have slow-growing populations and are at risk of complete extinction through hunting. There are some species of sharks with restricted distributions that are at risk of extinction. The groupers are another population of generally slow-growing fishes that, in the Caribbean, includes a number of species that are at risk of extinction from overfishing.
Coral reefs are extremely diverse marine ecosystems that face peril from several processes. Reefs are home to 1/3 of the world’s marine fish species—about 4,000 species—despite making up only 1 percent of marine habitat. Most home marine aquaria are stocked with wild-caught organisms, not cultured organisms. Although no species is known to have been driven extinct by the pet trade in marine species, there are studies showing that populations of some species have declined in response to harvesting, indicating that the harvest is not sustainable at those levels. There are concerns about the effect of the pet trade on some terrestrial species such as turtles, amphibians, birds, plants, and even the orangutan.
Bush meat is the generic term used for wild animals killed for food. Hunting is practiced throughout the world, but hunting practices, particularly in equatorial Africa and parts of Asia, are believed to threaten several species with extinction. Traditionally, bush meat in Africa was hunted to feed families directly; however, recent commercialization of the practice now has bush meat available in grocery stores, which has increased harvest rates to the level of unsustainability. Additionally, human population growth has increased the need for protein foods that are not being met from agriculture. Species threatened by the bush meat trade are mostly mammals including many primates living in the Congo basin.
Exotic Species
Exotic species are species that have been intentionally or unintentionally introduced by humans into an ecosystem in which they did not evolve. Such introductions likely occur frequently as natural phenomena. For example, Kudzu (Pueraria lobata), which is native to Japan, was introduced in the United States in 1876. It was later planted for soil conservation. Problematically, it grows too well in the southeastern United States—up to a foot a day. It is now a pest species and covers over 7 million acres in the southeastern United States. If an introduced species is able to survive in its new habitat, that introduction is now reflected in the observed range of the species. Human transportation of people and goods, including the intentional transport of organisms for trade, has dramatically increased the introduction of species into new ecosystems, sometimes at distances that are well beyond the capacity of the species to ever travel itself and outside the range of the species’ natural predators.
Most exotic species introductions probably fail because of the low number of individuals introduced or poor adaptation to the ecosystem they enter. Some species, however, possess preadaptations that can make them especially successful in a new ecosystem. These exotic species often undergo dramatic population increases in their new habitat and reset the ecological conditions in the new environment, threatening the species that exist there. For this reason, exotic species are also called invasive species. Exotic species can threaten other species through competition for resources, predation, or disease.
Link to Learning
Explore an interactive global database of exotic or invasive species.
Lakes and islands are particularly vulnerable to extinction threats from introduced species. In Lake Victoria, as mentioned earlier, the intentional introduction of the Nile perch was largely responsible for the extinction of about 200 species of cichlids. The accidental introduction of the brown tree snake via aircraft (Figure \(3\)) from the Solomon Islands to Guam in 1950 has led to the extinction of three species of birds and three to five species of reptiles endemic to the island. Several other species are still threatened. The brown tree snake is adept at exploiting human transportation as a means to migrate; one was even found on an aircraft arriving in Corpus Christi, Texas. Constant vigilance on the part of airport, military, and commercial aircraft personnel is required to prevent the snake from moving from Guam to other islands in the Pacific, especially Hawaii. Islands do not make up a large area of land on the globe, but they do contain a disproportionate number of endemic species because of their isolation from mainland ancestors.
It now appears that the global decline in amphibian species recognized in the 1990s is, in some part, caused by the fungus Batrachochytrium dendrobatidis, which causes the disease chytridiomycosis (Figure \(4\)). There is evidence that the fungus is native to Africa and may have been spread throughout the world by transport of a commonly used laboratory and pet species: the African clawed toad (Xenopus laevis). It may well be that biologists themselves are responsible for spreading this disease worldwide. The North American bullfrog, Rana catesbeiana, which has also been widely introduced as a food animal but which easily escapes captivity, survives most infections of Batrachochytrium dendrobatidis and can act as a reservoir for the disease.
Early evidence suggests that another fungal pathogen, Geomyces destructans, introduced from Europe is responsible for white-nose syndrome, which infects cave-hibernating bats in eastern North America and has spread from a point of origin in western New York State (Figure \(5\)). The disease has decimated bat populations and threatens extinction of species already listed as endangered: the Indiana bat, Myotis sodalis, and potentially the Virginia big-eared bat, Corynorhinus townsendii virginianus. How the fungus was introduced is unclear, but one logical presumption would be that recreational cavers unintentionally brought the fungus on clothes or equipment from Europe.
Climate Change
Climate change, and specifically the anthropogenic (meaning, caused by humans) warming trend presently underway, is recognized as a major extinction threat, particularly when combined with other threats such as habitat loss. Scientists disagree about the likely magnitude of the effects, with extinction rate estimates ranging from 15 percent to 40 percent of species committed to extinction by 2050. Scientists do agree, however, that climate change will alter regional climates, including rainfall and snowfall patterns, making habitats less hospitable to the species living in them. The warming trend will shift colder climates toward the north and south poles, forcing species to move with their adapted climate norms while facing habitat gaps along the way. The shifting ranges will impose new competitive regimes on species as they find themselves in contact with other species not present in their historic range. One such unexpected species contact is between polar bears and grizzly bears. Previously, these two species had separate ranges. Now, their ranges are overlapping and there are documented cases of these two species mating and producing viable offspring. Changing climates also throw off species’ delicate timing adaptations to seasonal food resources and breeding times. Many contemporary mismatches to shifts in resource availability and timing have already been documented.
Range shifts are already being observed: for example, some European bird species ranges have moved 91 km northward. The same study suggested that the optimal shift based on warming trends was double that distance, suggesting that the populations are not moving quickly enough. Range shifts have also been observed in plants, butterflies, other insects, freshwater fishes, reptiles, and mammals.
Climate gradients will also move up mountains, eventually crowding species higher in altitude and eliminating the habitat for those species adapted to the highest elevations. Some climates will completely disappear. The rate of warming appears to be accelerated in the arctic, which is recognized as a serious threat to polar bear populations that require sea ice to hunt seals during the winter months: seals are the only source of protein available to polar bears. A trend to decreasing sea ice coverage has occurred since observations began in the mid-twentieth century. The rate of decline observed in recent years is far greater than previously predicted by climate models.
Finally, global warming will raise ocean levels due to melt water from glaciers and the greater volume of warmer water. Shorelines will be inundated, reducing island size, which will have an effect on some species, and a number of islands will disappear entirely. Additionally, the gradual melting and subsequent refreezing of the poles, glaciers, and higher elevation mountains—a cycle that has provided freshwater to environments for centuries—will also be jeopardized. This could result in an overabundance of salt water and a shortage of fresh water.
Summary
The core threats to biodiversity are human population growth and unsustainable resource use. To date, the most significant causes of extinctions are habitat loss, introduction of exotic species, and overharvesting. Climate change is predicted to be a significant cause of extinctions in the coming century. Habitat loss occurs through deforestation, damming of rivers, and other activities. Overharvesting is a threat particularly to aquatic species, while the taking of bush meat in the humid tropics threatens many species in Asia, Africa, and the Americas. Exotic species have been the cause of a number of extinctions and are especially damaging to islands and lakes. Exotic species’ introductions are increasing because of the increased mobility of human populations and growing global trade and transportation. Climate change is forcing range changes that may lead to extinction. It is also affecting adaptations to the timing of resource availability that negatively affects species in seasonal environments. The impacts of climate change are greatest in the arctic. Global warming will also raise sea levels, eliminating some islands and reducing the area of all others.
Art Connections
Converting a prairie to a farm field is an example of ________.
1. overharvesting
2. habitat loss
3. exotic species
4. climate change
Answer
B
Glossary
bush meat
wild-caught animal used as food (typically mammals, birds, and reptiles); usually referring to hunting in the tropics of sub-Saharan Africa, Asia, and the Americas
chytridiomycosis
disease of amphibians caused by the fungus Batrachochytrium dendrobatidis; thought to be a major cause of the global amphibian decline
exotic species
(also, invasive species) species that has been introduced to an ecosystem in which it did not evolve
tragedy of the commons
economic principle that resources held in common will inevitably be overexploited
white-nose syndrome
disease of cave-hibernating bats in the eastern United States and Canada associated with the fungus Geomyces destructans
58.03: Factors Responsible for Extinction
Skills to Develop
• Identify new technologies for describing biodiversity
• Explain the legislative framework for conservation
• Describe principles and challenges of conservation preserve design
• Identify examples of the effects of habitat restoration
• Discuss the role of zoos in biodiversity conservation
Preserving biodiversity is an extraordinary challenge that must be met by greater understanding of biodiversity itself, changes in human behavior and beliefs, and various preservation strategies.
Measuring Biodiversity
The technology of molecular genetics and data processing and storage are maturing to the point where cataloguing the planet’s species in an accessible way is close to feasible. DNA barcoding is one molecular genetic method, which takes advantage of rapid evolution in a mitochondrial gene present in eukaryotes, excepting the plants, to identify species using the sequence of portions of the gene. Plants may be barcoded using a combination of chloroplast genes. Rapid mass sequencing machines make the molecular genetics portion of the work relatively inexpensive and quick. Computer resources store and make available the large volumes of data. Projects are currently underway to use DNA barcoding to catalog museum specimens, which have already been named and studied, as well as testing the method on less studied groups. As of mid 2012, close to 150,000 named species had been barcoded. Early studies suggest there are significant numbers of undescribed species that looked too much like sibling species to previously be recognized as different. These now can be identified with DNA barcoding.
Numerous computer databases now provide information about named species and a framework for adding new species. However, as already noted, at the present rate of description of new species, it will take close to 500 years before the complete catalog of life is known. Many, perhaps most, species on the planet do not have that much time.
There is also the problem of understanding which species known to science are threatened and to what degree they are threatened. This task is carried out by the non-profit IUCN which, as previously mentioned, maintains the Red List—an online listing of endangered species categorized by taxonomy, type of threat, and other criteria (Figure \(1\)). The Red List is supported by scientific research. In 2011, the list contained 61,000 species, all with supporting documentation.
Art Connection
Which of the following statements is not supported by this graph?
1. There are more vulnerable fishes than critically endangered and endangered fishes combined.
2. There are more critically endangered amphibians than vulnerable, endangered and critically endangered reptiles combined.
3. Within each group, there are more critically endangered species than vulnerable species.
4. A greater percentage of bird species are critically endangered than mollusk species.
Changing Human Behavior
Legislation throughout the world has been enacted to protect species. The legislation includes international treaties as well as national and state laws. The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) treaty came into force in 1975. The treaty, and the national legislation that supports it, provides a legal framework for preventing approximately 33,000 listed species from being transported across nations’ borders, thus protecting them from being caught or killed when international trade is involved. The treaty is limited in its reach because it only deals with international movement of organisms or their parts. It is also limited by various countries’ ability or willingness to enforce the treaty and supporting legislation. The illegal trade in organisms and their parts is probably a market in the hundreds of millions of dollars. Illegal wildlife trade is monitored by another non-profit: Trade Records Analysis of Flora and Fauna in Commerce (TRAFFIC).
Within many countries there are laws that protect endangered species and regulate hunting and fishing. In the United States, the Endangered Species Act (ESA) was enacted in 1973. Species at risk are listed by the Act; the U.S. Fish & Wildlife Service is required by law to develop management plans that protect the listed species and bring them back to sustainable numbers. The Act, and others like it in other countries, is a useful tool, but it suffers because it is often difficult to get a species listed, or to get an effective management plan in place once it is listed. Additionally, species may be controversially taken off the list without necessarily having had a change in their situation. More fundamentally, the approach to protecting individual species rather than entire ecosystems is both inefficient and focuses efforts on a few highly visible and often charismatic species, perhaps at the expense of other species that go unprotected. At the same time, the Act has a critical habitat provision outlined in the recovery mechanism that may benefit species other than the one targeted for management.
The Migratory Bird Treaty Act (MBTA) is an agreement between the United States and Canada that was signed into law in 1918 in response to declines in North American bird species caused by hunting. The Act now lists over 800 protected species. It makes it illegal to disturb or kill the protected species or distribute their parts (much of the hunting of birds in the past was for their feathers).
The international response to global warming has been mixed. The Kyoto Protocol, an international agreement that came out of the United Nations Framework Convention on Climate Change that committed countries to reducing greenhouse gas emissions by 2012, was ratified by some countries, but spurned by others. Two important countries in terms of their potential impact that did not ratify the Kyoto Protocol were the United States and China. The United States rejected it as a result of a powerful fossil fuel industry and China because of a concern it would stifle the nation’s growth. Some goals for reduction in greenhouse gasses were met and exceeded by individual countries, but worldwide, the effort to limit greenhouse gas production is not succeeding. The intended replacement for the Kyoto Protocol has not materialized because governments cannot agree on timelines and benchmarks. Meanwhile, climate scientists predict the resulting costs to human societies and biodiversity will be high.
As already mentioned, the private non-profit sector plays a large role in the conservation effort both in North America and around the world. The approaches range from species-specific organizations to the broadly focused IUCN and TRAFFIC. The Nature Conservancy takes a novel approach. It purchases land and protects it in an attempt to set up preserves for ecosystems. Ultimately, human behavior will change when human values change. At present, the growing urbanization of the human population is a force that poses challenges to the valuing of biodiversity.
Conservation in Preserves
Establishment of wildlife and ecosystem preserves is one of the key tools in conservation efforts. A preserve is an area of land set aside with varying degrees of protection for the organisms that exist within the boundaries of the preserve. Preserves can be effective in the short term for protecting both species and ecosystems, but they face challenges that scientists are still exploring to strengthen their viability as long-term solutions.
How Much Area to Preserve?
Due to the way protected lands are allocated (they tend to contain less economically valuable resources rather than being set aside specifically for the species or ecosystems at risk) and the way biodiversity is distributed, determining a target percentage of land or marine habitat that should be protected to maintain biodiversity levels is challenging. The IUCN World Parks Congress estimated that 11.5 percent of Earth’s land surface was covered by preserves of various kinds in 2003. This area is greater than previous goals; however, it only represents 9 out of 14 recognized major biomes. Research has shown that 12 percent of all species live only outside preserves; these percentages are much higher when only threatened species and high quality preserves are considered. For example, high quality preserves include only about 50 percent of threatened amphibian species. The conclusion must be that either the percentage of area protected must increase, or the percentage of high quality preserves must increase, or preserves must be targeted with greater attention to biodiversity protection. Researchers argue that more attention to the latter solution is required.
Preserve Design
There has been extensive research into optimal preserve designs for maintaining biodiversity. The fundamental principle behind much of the research has been the seminal theoretical work of Robert H. MacArthur and Edward O. Wilson published in 1967 on island biogeography.1 This work sought to understand the factors affecting biodiversity on islands. The fundamental conclusion was that biodiversity on an island was a function of the origin of species through migration, speciation, and extinction on that island. Islands farther from a mainland are harder to get to, so migration is lower and the equilibrium number of species is lower. Within island populations, evidence suggests that the number of species gradually increases to a level similar to the numbers on the mainland from which the species is suspected to have migrated. In addition, smaller islands are harder to find, so their immigration rates for new species are lower. Smaller islands are also less geographically diverse so there are fewer niches to promote speciation. And finally, smaller islands support smaller populations, so the probability of extinction is higher.
As islands get larger, the number of species accelerates, although the effect of island area on species numbers is not a direct correlation. Conservation preserves can be seen as “islands” of habitat within “an ocean” of non-habitat. For a species to persist in a preserve, the preserve must be large enough. The critical size depends, in part, on the home range that is characteristic of the species. A preserve for wolves, which range hundreds of kilometers, must be much larger than a preserve for butterflies, which might range within ten kilometers during its lifetime. But larger preserves have more core area of optimal habitat for individual species, they have more niches to support more species, and they attract more species because they can be found and reached more easily.
Preserves perform better when there are buffer zones around them of suboptimal habitat. The buffer allows organisms to exit the boundaries of the preserve without immediate negative consequences from predation or lack of resources. One large preserve is better than the same area of several smaller preserves because there is more core habitat unaffected by edges. For this same reason, preserves in the shape of a square or circle will be better than a preserve with many thin “arms.” If preserves must be smaller, then providing wildlife corridors between them so that individuals and their genes can move between the preserves, for example along rivers and streams, will make the smaller preserves behave more like a large one. All of these factors are taken into consideration when planning the nature of a preserve before the land is set aside.
In addition to the physical, biological, and ecological specifications of a preserve, there are a variety of policy, legislative, and enforcement specifications related to uses of the preserve for functions other than protection of species. These can include anything from timber extraction, mineral extraction, regulated hunting, human habitation, and nondestructive human recreation. Many of these policy decisions are made based on political pressures rather than conservation considerations. In some cases, wildlife protection policies have been so strict that subsistence-living indigenous populations have been forced from ancestral lands that fell within a preserve. In other cases, even if a preserve is designed to protect wildlife, if the protections are not or cannot be enforced, the preserve status will have little meaning in the face of illegal poaching and timber extraction. This is a widespread problem with preserves in areas of the tropics.
Limitations on Preserves
Some of the limitations on preserves as conservation tools are evident from the discussion of preserve design. Political and economic pressures typically make preserves smaller, never larger, so setting aside areas that are large enough is difficult. If the area set aside is sufficiently large, there may not be sufficient area to create a buffer around the preserve. In this case, an area on the outer edges of the preserve inevitably becomes a riskier suboptimal habitat for the species in the preserve. Enforcement of protections is also a significant issue in countries without the resources or political will to prevent poaching and illegal resource extraction.
Climate change will create inevitable problems with the location of preserves. The species within them will migrate to higher latitudes as the habitat of the preserve becomes less favorable. Scientists are planning for the effects of global warming on future preserves and striving to predict the need for new preserves to accommodate anticipated changes to habitats; however, the end effectiveness is tenuous since these efforts are prediction based.
Finally, an argument can be made that conservation preserves reinforce the cultural perception that humans are separate from nature, can exist outside of it, and can only operate in ways that do damage to biodiversity. Creating preserves reduces the pressure on human activities outside the preserves to be sustainable and non-damaging to biodiversity. Ultimately, the political, economic, and human demographic pressures will degrade and reduce the size of conservation preserves if the activities outside them are not altered to be less damaging to biodiversity.
Link to Learning
An interactive global data system of protected areas can be found at website. Review data about individual protected areas by location or study statistics on protected areas by country or region.
Habitat Restoration
Habitat restoration holds considerable promise as a mechanism for restoring and maintaining biodiversity. Of course once a species has become extinct, its restoration is impossible. However, restoration can improve the biodiversity of degraded ecosystems. Reintroducing wolves, a top predator, to Yellowstone National Park in 1995 led to dramatic changes in the ecosystem that increased biodiversity. The wolves (Figure \(2\)) function to suppress elk and coyote populations and provide more abundant resources to the guild of carrion eaters. Reducing elk populations has allowed revegetation of riparian areas, which has increased the diversity of species in that habitat. Decreasing the coyote population has increased the populations of species that were previously suppressed by this predator. The number of species of carrion eaters has increased because of the predatory activities of the wolves. In this habitat, the wolf is a keystone species, meaning a species that is instrumental in maintaining diversity in an ecosystem. Removing a keystone species from an ecological community may cause a collapse in diversity. The results from the Yellowstone experiment suggest that restoring a keystone species can have the effect of restoring biodiversity in the community. Ecologists have argued for the identification of keystone species where possible and for focusing protection efforts on those species; likewise, it also makes sense to attempt to return them to their ecosystem if they have been removed.
Other large-scale restoration experiments underway involve dam removal. In the United States, since the mid-1980s, many aging dams are being considered for removal rather than replacement because of shifting beliefs about the ecological value of free-flowing rivers and because many dams no longer provide the benefit and functions that they did when they were first built. The measured benefits of dam removal include restoration of naturally fluctuating water levels (the purpose of dams is frequently to reduce variation in river flows), which leads to increased fish diversity and improved water quality. In the Pacific Northwest, dam removal projects are expected to increase populations of salmon, which is considered a keystone species because it transports key nutrients to inland ecosystems during its annual spawning migrations. In other regions such as the Atlantic coast, dam removal has allowed the return of spawning anadromous fish species (species that are born in fresh water, live most of their lives in salt water, and return to fresh water to spawn). Some of the largest dam removal projects have yet to occur or have happened too recently for the consequences to be measured. The large-scale ecological experiments that these removal projects constitute will provide valuable data for other dam projects slated either for removal or construction.
The Role of Captive Breeding
Zoos have sought to play a role in conservation efforts both through captive breeding programs and education. The transformation of the missions of zoos from collection and exhibition facilities to organizations that are dedicated to conservation is ongoing. In general, it has been recognized that, except in some specific targeted cases, captive breeding programs for endangered species are inefficient and often prone to failure when the species are reintroduced to the wild. Zoo facilities are far too limited to contemplate captive breeding programs for the numbers of species that are now at risk. Education is another potential positive impact of zoos on conservation efforts, particularly given the global trend to urbanization and the consequent reduction in contacts between people and wildlife. A number of studies have been performed to look at the effectiveness of zoos on people’s attitudes and actions regarding conservation; at present, the results tend to be mixed.
Summary
New technological methods such as DNA barcoding and information processing and accessibility are facilitating the cataloging of the planet’s biodiversity. There is also a legislative framework for biodiversity protection. International treaties such as CITES regulate the transportation of endangered species across international borders. Legislation within individual countries protecting species and agreements on global warming have had limited success; there is at present no international agreement on targets for greenhouse gas emissions. In the United States, the Endangered Species Act protects listed species but is hampered by procedural difficulties and a focus on individual species. The Migratory Bird Act is an agreement between Canada and the United States to protect migratory birds. The non-profit sector is also very active in conservation efforts in a variety of ways.
Conservation preserves are a major tool in biodiversity protection. Presently, 11 percent of Earth’s land surface is protected in some way. The science of island biogeography has informed the optimal design of preserves; however, preserves have limitations imposed by political and economic forces. In addition, climate change will limit the effectiveness of preserves in the future. A downside of preserves is that they may lessen the pressure on human societies to function more sustainably outside the preserves.
Habitat restoration has the potential to restore ecosystems to previous biodiversity levels before species become extinct. Examples of restoration include reintroduction of keystone species and removal of dams on rivers. Zoos have attempted to take a more active role in conservation and can have a limited role in captive breeding programs. Zoos also may have a useful role in education.
Art Connections
Figure \(1\): Which of the following statements is not supported by this graph?
1. There are more vulnerable fishes than critically endangered and endangered fishes combined.
2. There are more critically endangered amphibians than vulnerable, endangered and critically endangered reptiles combined.
3. Within each group, there are more critically endangered species than vulnerable species.
4. A greater percentage of bird species are critically endangered than mollusk species.
Answer
C
Footnotes
1. 1 Robert H. MacArthur and Edward O. Wilson, E. O., The Theory of Island Biogeography (Princeton, N.J.: Princeton University Press, 1967).
Glossary
DNA barcoding
molecular genetic method for identifying a unique genetic sequence to associate with a species | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/58%3A_Conservation_Biology/58.03%3A_Factors_Responsible_for_Extinction/58.3.01%3A_Preserving_Biodiversity.txt |
Skills to Develop
• Identify new technologies for describing biodiversity
• Explain the legislative framework for conservation
• Describe principles and challenges of conservation preserve design
• Identify examples of the effects of habitat restoration
• Discuss the role of zoos in biodiversity conservation
Preserving biodiversity is an extraordinary challenge that must be met by greater understanding of biodiversity itself, changes in human behavior and beliefs, and various preservation strategies.
Measuring Biodiversity
The technology of molecular genetics and data processing and storage are maturing to the point where cataloguing the planet’s species in an accessible way is close to feasible. DNA barcoding is one molecular genetic method, which takes advantage of rapid evolution in a mitochondrial gene present in eukaryotes, excepting the plants, to identify species using the sequence of portions of the gene. Plants may be barcoded using a combination of chloroplast genes. Rapid mass sequencing machines make the molecular genetics portion of the work relatively inexpensive and quick. Computer resources store and make available the large volumes of data. Projects are currently underway to use DNA barcoding to catalog museum specimens, which have already been named and studied, as well as testing the method on less studied groups. As of mid 2012, close to 150,000 named species had been barcoded. Early studies suggest there are significant numbers of undescribed species that looked too much like sibling species to previously be recognized as different. These now can be identified with DNA barcoding.
Numerous computer databases now provide information about named species and a framework for adding new species. However, as already noted, at the present rate of description of new species, it will take close to 500 years before the complete catalog of life is known. Many, perhaps most, species on the planet do not have that much time.
There is also the problem of understanding which species known to science are threatened and to what degree they are threatened. This task is carried out by the non-profit IUCN which, as previously mentioned, maintains the Red List—an online listing of endangered species categorized by taxonomy, type of threat, and other criteria (Figure \(1\)). The Red List is supported by scientific research. In 2011, the list contained 61,000 species, all with supporting documentation.
Art Connection
Which of the following statements is not supported by this graph?
1. There are more vulnerable fishes than critically endangered and endangered fishes combined.
2. There are more critically endangered amphibians than vulnerable, endangered and critically endangered reptiles combined.
3. Within each group, there are more critically endangered species than vulnerable species.
4. A greater percentage of bird species are critically endangered than mollusk species.
Changing Human Behavior
Legislation throughout the world has been enacted to protect species. The legislation includes international treaties as well as national and state laws. The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) treaty came into force in 1975. The treaty, and the national legislation that supports it, provides a legal framework for preventing approximately 33,000 listed species from being transported across nations’ borders, thus protecting them from being caught or killed when international trade is involved. The treaty is limited in its reach because it only deals with international movement of organisms or their parts. It is also limited by various countries’ ability or willingness to enforce the treaty and supporting legislation. The illegal trade in organisms and their parts is probably a market in the hundreds of millions of dollars. Illegal wildlife trade is monitored by another non-profit: Trade Records Analysis of Flora and Fauna in Commerce (TRAFFIC).
Within many countries there are laws that protect endangered species and regulate hunting and fishing. In the United States, the Endangered Species Act (ESA) was enacted in 1973. Species at risk are listed by the Act; the U.S. Fish & Wildlife Service is required by law to develop management plans that protect the listed species and bring them back to sustainable numbers. The Act, and others like it in other countries, is a useful tool, but it suffers because it is often difficult to get a species listed, or to get an effective management plan in place once it is listed. Additionally, species may be controversially taken off the list without necessarily having had a change in their situation. More fundamentally, the approach to protecting individual species rather than entire ecosystems is both inefficient and focuses efforts on a few highly visible and often charismatic species, perhaps at the expense of other species that go unprotected. At the same time, the Act has a critical habitat provision outlined in the recovery mechanism that may benefit species other than the one targeted for management.
The Migratory Bird Treaty Act (MBTA) is an agreement between the United States and Canada that was signed into law in 1918 in response to declines in North American bird species caused by hunting. The Act now lists over 800 protected species. It makes it illegal to disturb or kill the protected species or distribute their parts (much of the hunting of birds in the past was for their feathers).
The international response to global warming has been mixed. The Kyoto Protocol, an international agreement that came out of the United Nations Framework Convention on Climate Change that committed countries to reducing greenhouse gas emissions by 2012, was ratified by some countries, but spurned by others. Two important countries in terms of their potential impact that did not ratify the Kyoto Protocol were the United States and China. The United States rejected it as a result of a powerful fossil fuel industry and China because of a concern it would stifle the nation’s growth. Some goals for reduction in greenhouse gasses were met and exceeded by individual countries, but worldwide, the effort to limit greenhouse gas production is not succeeding. The intended replacement for the Kyoto Protocol has not materialized because governments cannot agree on timelines and benchmarks. Meanwhile, climate scientists predict the resulting costs to human societies and biodiversity will be high.
As already mentioned, the private non-profit sector plays a large role in the conservation effort both in North America and around the world. The approaches range from species-specific organizations to the broadly focused IUCN and TRAFFIC. The Nature Conservancy takes a novel approach. It purchases land and protects it in an attempt to set up preserves for ecosystems. Ultimately, human behavior will change when human values change. At present, the growing urbanization of the human population is a force that poses challenges to the valuing of biodiversity.
Conservation in Preserves
Establishment of wildlife and ecosystem preserves is one of the key tools in conservation efforts. A preserve is an area of land set aside with varying degrees of protection for the organisms that exist within the boundaries of the preserve. Preserves can be effective in the short term for protecting both species and ecosystems, but they face challenges that scientists are still exploring to strengthen their viability as long-term solutions.
How Much Area to Preserve?
Due to the way protected lands are allocated (they tend to contain less economically valuable resources rather than being set aside specifically for the species or ecosystems at risk) and the way biodiversity is distributed, determining a target percentage of land or marine habitat that should be protected to maintain biodiversity levels is challenging. The IUCN World Parks Congress estimated that 11.5 percent of Earth’s land surface was covered by preserves of various kinds in 2003. This area is greater than previous goals; however, it only represents 9 out of 14 recognized major biomes. Research has shown that 12 percent of all species live only outside preserves; these percentages are much higher when only threatened species and high quality preserves are considered. For example, high quality preserves include only about 50 percent of threatened amphibian species. The conclusion must be that either the percentage of area protected must increase, or the percentage of high quality preserves must increase, or preserves must be targeted with greater attention to biodiversity protection. Researchers argue that more attention to the latter solution is required.
Preserve Design
There has been extensive research into optimal preserve designs for maintaining biodiversity. The fundamental principle behind much of the research has been the seminal theoretical work of Robert H. MacArthur and Edward O. Wilson published in 1967 on island biogeography.1 This work sought to understand the factors affecting biodiversity on islands. The fundamental conclusion was that biodiversity on an island was a function of the origin of species through migration, speciation, and extinction on that island. Islands farther from a mainland are harder to get to, so migration is lower and the equilibrium number of species is lower. Within island populations, evidence suggests that the number of species gradually increases to a level similar to the numbers on the mainland from which the species is suspected to have migrated. In addition, smaller islands are harder to find, so their immigration rates for new species are lower. Smaller islands are also less geographically diverse so there are fewer niches to promote speciation. And finally, smaller islands support smaller populations, so the probability of extinction is higher.
As islands get larger, the number of species accelerates, although the effect of island area on species numbers is not a direct correlation. Conservation preserves can be seen as “islands” of habitat within “an ocean” of non-habitat. For a species to persist in a preserve, the preserve must be large enough. The critical size depends, in part, on the home range that is characteristic of the species. A preserve for wolves, which range hundreds of kilometers, must be much larger than a preserve for butterflies, which might range within ten kilometers during its lifetime. But larger preserves have more core area of optimal habitat for individual species, they have more niches to support more species, and they attract more species because they can be found and reached more easily.
Preserves perform better when there are buffer zones around them of suboptimal habitat. The buffer allows organisms to exit the boundaries of the preserve without immediate negative consequences from predation or lack of resources. One large preserve is better than the same area of several smaller preserves because there is more core habitat unaffected by edges. For this same reason, preserves in the shape of a square or circle will be better than a preserve with many thin “arms.” If preserves must be smaller, then providing wildlife corridors between them so that individuals and their genes can move between the preserves, for example along rivers and streams, will make the smaller preserves behave more like a large one. All of these factors are taken into consideration when planning the nature of a preserve before the land is set aside.
In addition to the physical, biological, and ecological specifications of a preserve, there are a variety of policy, legislative, and enforcement specifications related to uses of the preserve for functions other than protection of species. These can include anything from timber extraction, mineral extraction, regulated hunting, human habitation, and nondestructive human recreation. Many of these policy decisions are made based on political pressures rather than conservation considerations. In some cases, wildlife protection policies have been so strict that subsistence-living indigenous populations have been forced from ancestral lands that fell within a preserve. In other cases, even if a preserve is designed to protect wildlife, if the protections are not or cannot be enforced, the preserve status will have little meaning in the face of illegal poaching and timber extraction. This is a widespread problem with preserves in areas of the tropics.
Limitations on Preserves
Some of the limitations on preserves as conservation tools are evident from the discussion of preserve design. Political and economic pressures typically make preserves smaller, never larger, so setting aside areas that are large enough is difficult. If the area set aside is sufficiently large, there may not be sufficient area to create a buffer around the preserve. In this case, an area on the outer edges of the preserve inevitably becomes a riskier suboptimal habitat for the species in the preserve. Enforcement of protections is also a significant issue in countries without the resources or political will to prevent poaching and illegal resource extraction.
Climate change will create inevitable problems with the location of preserves. The species within them will migrate to higher latitudes as the habitat of the preserve becomes less favorable. Scientists are planning for the effects of global warming on future preserves and striving to predict the need for new preserves to accommodate anticipated changes to habitats; however, the end effectiveness is tenuous since these efforts are prediction based.
Finally, an argument can be made that conservation preserves reinforce the cultural perception that humans are separate from nature, can exist outside of it, and can only operate in ways that do damage to biodiversity. Creating preserves reduces the pressure on human activities outside the preserves to be sustainable and non-damaging to biodiversity. Ultimately, the political, economic, and human demographic pressures will degrade and reduce the size of conservation preserves if the activities outside them are not altered to be less damaging to biodiversity.
Link to Learning
An interactive global data system of protected areas can be found at website. Review data about individual protected areas by location or study statistics on protected areas by country or region.
Habitat Restoration
Habitat restoration holds considerable promise as a mechanism for restoring and maintaining biodiversity. Of course once a species has become extinct, its restoration is impossible. However, restoration can improve the biodiversity of degraded ecosystems. Reintroducing wolves, a top predator, to Yellowstone National Park in 1995 led to dramatic changes in the ecosystem that increased biodiversity. The wolves (Figure \(2\)) function to suppress elk and coyote populations and provide more abundant resources to the guild of carrion eaters. Reducing elk populations has allowed revegetation of riparian areas, which has increased the diversity of species in that habitat. Decreasing the coyote population has increased the populations of species that were previously suppressed by this predator. The number of species of carrion eaters has increased because of the predatory activities of the wolves. In this habitat, the wolf is a keystone species, meaning a species that is instrumental in maintaining diversity in an ecosystem. Removing a keystone species from an ecological community may cause a collapse in diversity. The results from the Yellowstone experiment suggest that restoring a keystone species can have the effect of restoring biodiversity in the community. Ecologists have argued for the identification of keystone species where possible and for focusing protection efforts on those species; likewise, it also makes sense to attempt to return them to their ecosystem if they have been removed.
Other large-scale restoration experiments underway involve dam removal. In the United States, since the mid-1980s, many aging dams are being considered for removal rather than replacement because of shifting beliefs about the ecological value of free-flowing rivers and because many dams no longer provide the benefit and functions that they did when they were first built. The measured benefits of dam removal include restoration of naturally fluctuating water levels (the purpose of dams is frequently to reduce variation in river flows), which leads to increased fish diversity and improved water quality. In the Pacific Northwest, dam removal projects are expected to increase populations of salmon, which is considered a keystone species because it transports key nutrients to inland ecosystems during its annual spawning migrations. In other regions such as the Atlantic coast, dam removal has allowed the return of spawning anadromous fish species (species that are born in fresh water, live most of their lives in salt water, and return to fresh water to spawn). Some of the largest dam removal projects have yet to occur or have happened too recently for the consequences to be measured. The large-scale ecological experiments that these removal projects constitute will provide valuable data for other dam projects slated either for removal or construction.
The Role of Captive Breeding
Zoos have sought to play a role in conservation efforts both through captive breeding programs and education. The transformation of the missions of zoos from collection and exhibition facilities to organizations that are dedicated to conservation is ongoing. In general, it has been recognized that, except in some specific targeted cases, captive breeding programs for endangered species are inefficient and often prone to failure when the species are reintroduced to the wild. Zoo facilities are far too limited to contemplate captive breeding programs for the numbers of species that are now at risk. Education is another potential positive impact of zoos on conservation efforts, particularly given the global trend to urbanization and the consequent reduction in contacts between people and wildlife. A number of studies have been performed to look at the effectiveness of zoos on people’s attitudes and actions regarding conservation; at present, the results tend to be mixed.
Summary
New technological methods such as DNA barcoding and information processing and accessibility are facilitating the cataloging of the planet’s biodiversity. There is also a legislative framework for biodiversity protection. International treaties such as CITES regulate the transportation of endangered species across international borders. Legislation within individual countries protecting species and agreements on global warming have had limited success; there is at present no international agreement on targets for greenhouse gas emissions. In the United States, the Endangered Species Act protects listed species but is hampered by procedural difficulties and a focus on individual species. The Migratory Bird Act is an agreement between Canada and the United States to protect migratory birds. The non-profit sector is also very active in conservation efforts in a variety of ways.
Conservation preserves are a major tool in biodiversity protection. Presently, 11 percent of Earth’s land surface is protected in some way. The science of island biogeography has informed the optimal design of preserves; however, preserves have limitations imposed by political and economic forces. In addition, climate change will limit the effectiveness of preserves in the future. A downside of preserves is that they may lessen the pressure on human societies to function more sustainably outside the preserves.
Habitat restoration has the potential to restore ecosystems to previous biodiversity levels before species become extinct. Examples of restoration include reintroduction of keystone species and removal of dams on rivers. Zoos have attempted to take a more active role in conservation and can have a limited role in captive breeding programs. Zoos also may have a useful role in education.
Art Connections
Figure \(1\): Which of the following statements is not supported by this graph?
1. There are more vulnerable fishes than critically endangered and endangered fishes combined.
2. There are more critically endangered amphibians than vulnerable, endangered and critically endangered reptiles combined.
3. Within each group, there are more critically endangered species than vulnerable species.
4. A greater percentage of bird species are critically endangered than mollusk species.
Answer
C
Footnotes
1. 1 Robert H. MacArthur and Edward O. Wilson, E. O., The Theory of Island Biogeography (Princeton, N.J.: Princeton University Press, 1967).
Glossary
DNA barcoding
molecular genetic method for identifying a unique genetic sequence to associate with a species | textbooks/bio/Introductory_and_General_Biology/Map%3A_Raven_Biology_12th_Edition/58%3A_Conservation_Biology/58.05%3A_Approaches_for_Preserving_Endangered_Species_and_Ecosystems.txt |
Learning Objectives
Course Outcomes for this section:
Apply the scientific method to biological questions by designing experiments and using the resulting data to form a conclusion.
1. Design a controlled experiment to answer a biological question.
2. Predict the outcome of an experiment.
3. Collect, manipulate, and analyze quantitative and qualitative data
4. Answer a biological question using data.
Select, evaluate, and utilize discipline-specific information and literature to research a biological topic.
1. Differentiate between questions that can and cannot be answered using science.
2. Identify appropriate credible sources of information to research a topic.
3. Evaluate sources of information for their strengths and weaknesses.
4. Differentiate between popular and scholarly sources.
Like geology, physics, and chemistry, biology is a science that gathers knowledge about the natural world. Specifically, biology is the study of life. The discoveries of biology are made by a community of researchers who work individually and together using agreed-on methods. In this sense, biology, like all sciences is a social enterprise like politics or the arts. The methods of science include careful observation, record keeping, logical and mathematical reasoning, experimentation, and submitting conclusions to the scrutiny of others. Science also requires considerable imagination and creativity; a well-designed experiment is commonly described as elegant, or beautiful. Like politics, science has considerable practical implications and some science is dedicated to practical applications, such as the prevention of disease (see Figure \(1\)). Other science proceeds largely motivated by curiosity. Whatever its goal, there is no doubt that science, including biology, has transformed human existence and will continue to do so.
01: The Process of Science
Biology is a science, but what exactly is science? What does the study of biology share with other scientific disciplines? Science (from the Latin scientia, meaning “knowledge”) can be defined as knowledge about the natural world. Science is a very specific way of learning, or knowing, about the world. The history of the past 500 years demonstrates that science is a very powerful way of knowing about the world; it is largely responsible for the technological revolutions that have taken place during this time. There are however, areas of knowledge and human experience that the methods of science cannot be applied to. These include such things as answering purely moral questions, aesthetic questions, or what can be generally categorized as spiritual questions. Science can not investigate these areas because they are outside the realm of material phenomena, the phenomena of matter and energy, and can not be observed and measured.
The scientific method is a method of research with defined steps that include experiments and careful observation. One of the most important aspects of this method is the testing of hypotheses. A hypothesis is a suggested explanation for an event, which can be tested. Hypotheses, or tentative explanations, are generally produced within the context of a scientific theory. A scientific theory is a generally accepted, thoroughly tested and confirmed explanation for a set of observations or phenomena. Scientific theory is the foundation of scientific knowledge. In addition, in many scientific disciplines (less so in biology) there are scientific laws, often expressed in mathematical formulas, which describe how elements of nature will behave under certain specific conditions. There is not an evolution of hypotheses through theories to laws as if they represented some increase in certainty about the world. Hypotheses are the day-to-day material that scientists work with and they are developed within the context of theories. Laws are concise descriptions of parts of the world that are amenable to formulaic or mathematical description.
Query \(1\)
The scientific community has been debating for the last few decades about the value of different types of science. Is it valuable to pursue science for the sake of simply gaining knowledge, or does scientific knowledge only have worth if we can apply it to solving a specific problem or bettering our lives? This question focuses on the differences between two types of science: basic science and applied science.
• Basic science or “pure” science seeks to expand knowledge regardless of the short-term application of that knowledge. It is not focused on developing a product or a service of immediate public or commercial value. The immediate goal of basic science is knowledge for knowledge’s sake, though this does not mean that in the end it may not result in an application.
• In contrast, applied science or “technology,” aims to use science to solve real-world problems, making it possible, for example, to improve a crop yield, find a cure for a particular disease, or save animals threatened by a natural disaster. In applied science, the problem is usually defined for the researcher.
Some individuals may perceive applied science as “useful” and basic science as “useless.” A question these people might pose to a scientist advocating knowledge acquisition would be, “What for?” A careful look at the history of science, however, reveals that basic knowledge has resulted in many remarkable applications of great value. Many scientists think that a basic understanding of science is necessary before an application is developed; therefore, applied science relies on the results generated through basic science. Other scientists think that it is time to move on from basic science and instead to find solutions to actual problems. Both approaches are valid. It is true that there are problems that demand immediate attention; however, few solutions would be found without the help of the knowledge generated through basic science.
One example of how basic and applied science can work together to solve practical problems occurred after the discovery of DNA structure led to an understanding of the molecular mechanisms governing DNA replication. Strands of DNA, unique in every human, are found in our cells, where they provide the instructions necessary for life. During DNA replication, new copies of DNA are made, shortly before a cell divides to form new cells. Understanding the mechanisms of DNA replication enabled scientists to develop laboratory techniques that are now used to identify genetic diseases, pinpoint individuals who were at a crime scene, and determine paternity. Without basic science, it is unlikely that applied science would exist.
Another example of the link between basic and applied research is the Human Genome Project, a study in which each human chromosome was analyzed and mapped to determine the precise sequence of DNA subunits and the exact location of each gene. (The gene is the basic unit of heredity; an individual’s complete collection of genes is his or her genome.) Other organisms have also been studied as part of this project to gain a better understanding of human chromosomes. The Human Genome Project (Figure \(1\)) relied on basic research carried out with non-human organisms and, later, with the human genome. An important end goal eventually became using the data for applied research seeking cures for genetically related diseases.
While research efforts in both basic science and applied science are usually carefully planned, it is important to note that some discoveries are made by serendipity, that is, by means of a fortunate accident or a lucky surprise. Penicillin was discovered when biologist Alexander Fleming accidentally left a petri dish of Staphylococcus bacteria open. An unwanted mold grew, killing the bacteria. The mold turned out to be Penicillium, and a new antibiotic was discovered. Even in the highly organized world of science, luck—when combined with an observant, curious mind—can lead to unexpected breakthroughs. | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/01%3A_The_Process_of_Science/1.01%3A_The_Nature_of_Science.txt |
Biologists study the living world by posing questions about it and seeking science-based responses. This approach is common to other sciences as well and is often referred to as the scientific method. The scientific process was used even in ancient times, but it was first documented by England’s Sir Francis Bacon (1561–1626) (Figure \(1\)), who set up inductive methods for scientific inquiry. The scientific method is not exclusively used by biologists but can be applied to almost anything as a logical problem solving method.
Question
The scientific process typically starts with an observation (often a problem to be solved) that leads to a question. Remember that science is very good at answering questions having to do with observations about the natural world, but is very bad at answering questions having to do with morals, ethics, or personal opinions.
Questions that can be answered using science Questions that cannot be answered using science
• What is the optimum temperature for the growth of E. coli bacteria? • How tall is Santa Claus?
• Do birds prefer bird feeders of a specific color? • Do angels exist?
• What is the cause of this disease? • Which is better: classical music or rock and roll?
• How effective is this drug in treating this disease? • What are the ethical implications of human cloning?
Let’s think about a simple problem that starts with an observation and apply the scientific method to solve the problem. Imagine that one morning when you wake up and flip the switch to turn on your bedside lamp, the light won’t turn on. That is an observation that also describes a problem: the lights won’t turn on. Of course, you would next ask the question: “Why won’t the light turn on?”
Hypothesis
Recall that a hypothesis is a suggested explanation that can be tested. A hypothesis is NOT the question you are trying to answer – it is what you think the answer to the question will be and why. To solve a problem, several hypotheses may be proposed. For example, one hypothesis might be, “The light won’t turn on because the bulb is burned out.” But there could be other answers to the question, and therefore other hypotheses may be proposed. A second hypothesis might be, “The light won’t turn on because the lamp is unplugged” or “The light won’t turn on because the power is out.” A hypothesis should be based on credible background information. A hypothesis is NOT just a guess (not even an educated one), although it can be based on your prior experience (such as in the example where the light won’t turn on). In general, hypotheses in biology should be based on a credible, referenced source of information.
A hypothesis must be testable to ensure that it is valid. For example, a hypothesis that depends on what a dog thinks is not testable, because we can’t tell what a dog thinks. It should also be falsifiable, meaning that it can be disproven by experimental results. An example of an unfalsifiable hypothesis is “Red is a better color than blue.” There is no experiment that might show this statement to be false. To test a hypothesis, a researcher will conduct one or more experiments designed to eliminate one or more of the hypotheses. This is important: a hypothesis can be disproven, or eliminated, but it can never be proven. Science does not deal in proofs like mathematics. If an experiment fails to disprove a hypothesis, then that explanation (the hypothesis) is supported as the answer to the question. However, that doesn’t mean that later on, we won’t find a better explanation or design a better experiment that will be found to falsify the first hypothesis and lead to a better one.
Variables
A variable is any part of the experiment that can vary or change during the experiment. Typically, an experiment only tests one variable and all the other conditions in the experiment are held constant.
• The variable that is tested is known as the independent variable.
• The dependent variable is the thing (or things) that you are measuring as the outcome of your experiment.
• constant is a condition that is the same between all of the tested groups.
• A confounding variable is a condition that is not held constant that could affect the experimental results.
A hypothesis often has the format “If [I change the independent variable in this way] then [I will observe that the dependent variable does this] because [of some reason].” For example, the first hypothesis might be, “If you change the light bulb, then the light will turn on because the bulb is burned out.” In this experiment, the independent variable (the thing that you are testing) would be changing the light bulb and the dependent variable is whether or not the light turns on. It would be important to hold all the other aspects of the environment constant, for example not messing with the lamp cord or trying to turn the lamp on using a different light switch. If the entire house had lost power during the experiment because a car hit the power pole, that would be a confounding variable.
You may have learned that a hypothesis can be phrased as an “If..then…” statement. Simple hypotheses can be phrased that way (but they must also include a “because”), but more complicated hypotheses may require several sentences. It is also very easy to get confused by trying to put your hypothesis into this format. Hypotheses do not have to be phrased as “if..then..” statements, it is just sometimes a useful format.
Query \(1\)
Query \(2\)
Query \(3\)
Results
The results of your experiment are the data that you collect as the outcome. In the light experiment, your results are either that the light turns on or the light doesn’t turn on. Based on your results, you can make a conclusion. Your conclusion uses the results to answer your original question.
We can put the experiment with the light that won’t go in into the figure above:
1. Observation: the light won’t turn on.
2. Question: why won’t the light turn on?
3. Hypothesis: the lightbulb is burned out.
4. Prediction: if I change the lightbulb (independent variable), then the light will turn on (dependent variable).
5. Experiment: change the lightbulb while leaving all other variables the same.
6. Analyze the results: the light didn’t turn on.
7. Conclusion: The lightbulb isn’t burned out. The results do not support the hypothesis, time to develop a new one!
8. Hypothesis 2: the lamp is unplugged.
9. Prediction 2: if I plug in the lamp, then the light will turn on.
10. Experiment: plug in the lamp
11. Analyze the results: the light turned on!
12. Conclusion: The light wouldn’t turn on because the lamp was unplugged. The results support the hypothesis, it’s time to move on to the next experiment!
In practice, the scientific method is not as rigid and structured as it might at first appear. Sometimes an experiment leads to conclusions that favor a change in approach; often, an experiment brings entirely new scientific questions to the puzzle. Many times, science does not operate in a linear fashion; instead, scientists continually draw inferences and make generalizations, finding patterns as their research proceeds. Scientific reasoning is more complex than the scientific method alone suggests.
Control Groups
Another important aspect of designing an experiment is the presence of one or more control groups. A control group allows you to make a comparison that is important for interpreting your results. Control groups are samples that help you to determine that differences between your experimental groups are due to your treatment rather than a different variable – they eliminate alternate explanations for your results (including experimental error and experimenter bias). They increase reliability, often through the comparison of control measurements and measurements of the experimental groups. Often, the control group is a sample that is not treated with the independent variable, but is otherwise treated the same way as your experimental sample. This type of control group contains every feature of the experimental group except it is not given the manipulation that is hypothesized about (it does not get treated with the independent variable). Therefore, if the results of the experimental group differ from the control group, the difference must be due to the hypothesized manipulation, rather than some outside factor. It is common in complex experiments (such as those published in scientific journals) to have more control groups than experimental groups.
Example \(1\)
Question: Which fertilizer will produce the greatest number of tomatoes when applied to the plants?
Prediction and Hypothesis: If I apply different brands of fertilizer to tomato plants, the most tomatoes will be produced from plants watered with Brand A because Brand A advertises that it produces twice as many tomatoes as other leading brands.
Experiment: Purchase 10 tomato plants of the same type from the same nursery. Pick plants that are similar in size and age. Divide the plants into two groups of 5. Apply Brand A to the first group and Brand B to the second group according to the instructions on the packages. After 10 weeks, count the number of tomatoes on each plant.
Independent Variable: Brand of fertilizer.
Dependent Variable: Number of tomatoes.
The number of tomatoes produced depends on the brand of fertilizer applied to the plants.
Constants: amount of water, type of soil, size of pot, amount of light, type of tomato plant, length of time plants were grown.
Confounding variables: any of the above that are not held constant, plant health, diseases present in the soil or plant before it was purchased.
Results: Tomatoes fertilized with Brand A produced an average of 20 tomatoes per plant, while tomatoes fertilized with Brand B produced an average of 10 tomatoes per plant.
You’d want to use Brand A next time you grow tomatoes, right? But what if I told you that plants grown without fertilizer produced an average of 30 tomatoes per plant! Now what will you use on your tomatoes?
Results including control group: Tomatoes which received no fertilizer produced more tomatoes than either brand of fertilizer.
Conclusion: Although Brand A fertilizer produced more tomatoes than Brand B, neither fertilizer should be used because plants grown without fertilizer produced the most tomatoes!
Positive control groups are often used to show that the experiment is valid and that everything has worked correctly. You can think of a positive control group as being a group where you should be able to observe the thing that you are measuring (“the thing” should happen). The conditions in a positive control group should guarantee a positive result. If the positive control group doesn’t work, there may be something wrong with the experimental procedure.
Negative control groups are used to show whether a treatment had any effect. If your treated sample is the same as your negative control group, your treatment had no effect. You can also think of a negative control group as being a group where you should NOT be able to observe the thing that you are measuring (“the thing” shouldn’t happen), or where you should not observe any change in the thing that you are measuring (there is no difference between the treated and control group). The conditions in a negative control group should guarantee a negative result. A placebo group is an example of a negative control group.
As a general rule, you need a positive control to validate a negative result, and a negative control to validate a positive result.
• You read an article in the NY Times that says some spinach is contaminated with Salmonella. You want to test the spinach you have at home in your fridge, so you wet a sterile swab and wipe it on the spinach, then wipe the swab on a nutrient plate (petri plate).
• You observe growth. Does this mean that your spinach is really contaminated? Consider an alternate explanation for growth: the swab, the water, or the plate is contaminated with bacteria. You could use a negative control to determine which explanation is true. If a swab is wet and wiped on a nutrient plate, do bacteria grow?
• You don’t observe growth. Does this mean that your spinach is really safe? Consider an alternate explanation for no growth: Salmonella isn’t able to grow on the type of nutrient you used in your plates. You could use a positive control to determine which explanation is true. If you wipe a known sample of Salmonella bacteria on the plate, do bacteria grow?
• In a drug trial, one group of subjects are given a new drug, while a second group is given a placebo drug (a sugar pill; something which appears like the drug, but doesn’t contain the active ingredient). Reduction in disease symptoms are measured. The second group receiving the placebo is a negative control group. You might expect a reduction in disease symptoms purely because the person knows they are taking a drug so they should be getting better. If the group treated with the real drug does not show more a reduction in disease symptoms than the placebo group, the drug doesn’t really work. The placebo group sets a baseline against which the experimental group (treated with the drug) can be compared. A positive control group is not required for this experiment.
• In an experiment measuring the preference of birds for various types of food, a negative control group would be a “placebo feeder”. This would be the same type of feeder, but with no food in it. Birds might visit a feeder just because they are interested in it; an empty feeder would give a baseline level for bird visits. A positive control group might be a food that squirrels are known to like. This would be useful because if no squirrels visited any of the feeders, you couldn’t tell if this was because there were no squirrels around or because they didn’t like any of your food offerings!
• To test the effect of pH on the function of an enzyme, you would want a positive control group where you knew the enzyme would function (pH not changed) and a negative control group where you knew the enzyme would not function (no enzyme added). You need the positive control group so you know your enzyme is working: if you didn’t see a reaction in any of the tubes with the pH adjusted, you wouldn’t know if it was because the enzyme wasn’t working at all or because the enzyme just didn’t work at any of your tested pH. You need the negative control group so you can ensure that there is no reaction taking place in the absence of enzyme: if the reaction proceeds without the enzyme, your results are meaningless. | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/01%3A_The_Process_of_Science/1.02%3A_The_Scientific_Process.txt |
Types of Data
There are different types of data that can be collected in an experiment. Typically, we try to design experiments that collect objective, quantitative data.
Objective data is fact-based, measurable, and observable. This means that if two people made the same measurement with the same tool, they would get the same answer. The measurement is determined by the object that is being measured. The length of a worm measured with a ruler is an objective measurement. The observation that a chemical reaction in a test tube changed color is an objective measurement. Both of these are observable facts.
Subjective data is based on opinions, points of view, or emotional judgment. Subjective data might give two different answers when collected by two different people. The measurement is determined by the subject who is doing the measuring. Surveying people about which of two chemicals smells worse is a subjective measurement. Grading the quality of a presentation is a subjective measurement. Rating your relative happiness on a scale of 1-5 is a subjective measurement. All of these depend on the person who is making the observation – someone else might make these measurements differently.
Quantitative measurements gather numerical data. For example, measuring a worm as being 5cm in length is a quantitative measurement.
Qualitative measurements describe a quality, rather than a numerical value. Saying that one worm is longer than another worm is a qualitative measurement.
Quantitative Qualitative
Objective The chemical reaction has produced 5cm of bubbles. The chemical reaction has produced a lot of bubbles.
Subjective I give the amount of bubbles a score of 7 on a scale of 1-10. I think the bubbles are pretty.
After you have collected data in an experiment, you need to figure out the best way to present that data in a meaningful way. Depending on the type of data, and the story that you are trying to tell using that data, you may present your data in different ways.
Query \(1\)
Query \(2\)
Data Tables
The easiest way to organize data is by putting it into a data table. In most data tables, the independent variable (the variable that you are testing or changing on purpose) will be in the column to the left and the dependent variable(s) will be across the top of the table.
Be sure to:
• Label each row and column so that the table can be interpreted
• Include the units that are being used
• Add a descriptive caption for the table
Example \(1\)
You are evaluating the effect of different types of fertilizers on plant growth. You plant 12 tomato plants and divide them into three groups, where each group contains four plants. To the first group, you do not add fertilizer and the plants are watered with plain water. The second and third groups are watered with two different brands of fertilizer. After three weeks, you measure the growth of each plant in centimeters and calculate the average growth for each type of fertilizer.
The effect of different brands of fertilizer on tomato plant growth over three weeks
Treatment Plant Number
1 2 3 4 Average
No treatment 10 12 8 9 9.75
Brand A 15 16 14 12 14.25
Brand B 22 25 21 27 23.75
Scientific Method Review: Can you identify the key parts of the scientific method from this experiment?
• Independent variable – Type of treatment (brand of fertilizer)
• Dependent variable – plant growth in cm
• Control group(s) – Plants treated with no fertilizer
• Experimental group(s) – Plants treated with different brands of fertilizer
Graphing data
Graphs are used to display data because it is easier to see trends in the data when it is displayed visually compared to when it is displayed numerically in a table. Complicated data can often be displayed and interpreted more easily in a graph format than in a data table.
In a graph, the X-axis runs horizontally (side to side) and the Y-axis runs vertically (up and down). Typically, the independent variable will be shown on the X axis and the dependent variable will be shown on the Y axis (just like you learned in math class!).
Line Graph
Line graphs are the best type of graph to use when you are displaying a change in something over a continuous range. For example, you could use a line graph to display a change in temperature over time. Time is a continuous variable because it can have any value between two given measurements. It is measured along a continuum. Between 1 minute and 2 minutes are an infinite number of values, such as 1.1 minute or 1.93456 minutes.
Changes in several different samples can be shown on the same graph by using lines that differ in color, symbol, etc.
Bar Graph
Bar graphs are used to compare measurements between different groups. Bar graphs should be used when your data is not continuous, but rather is divided into different categories. If you counted the number of birds of different species, each species of bird would be its own category. There is no value between “robin” and “eagle”, so this data is not continuous.
Scatter Plot
Scatter plots are used to evaluate the relationship between two different continuous variables. These graphs compare changes in two different variables at once. For example, you could look at the relationship between height and weight. Both height and weight are continuous variables. You could not use a scatter plot to look at the relationship between number of children in a family and weight of each child because the number of children in a family is not a continuous variable: you can’t have 2.3 children in a family.
Query \(3\)
How to make a graph
1. Identify your independent and dependent variables.
2. Choose the correct type of graph by determining whether each variable is continuous or not.
3. Determine the values that are going to go on the X and Y axis. If the values are continuous, they need to be evenly spaced based on the value.
4. Label the X and Y axis, including units.
5. Graph your data.
6. Add a descriptive caption to your graph. Note that data tables are titled above the figure and graphs are captioned below the figure.
Example \(2\)
Let’s go back to the data from our fertilizer experiment and use it to make a graph. I’ve decided to graph only the average growth for the four plants because that is the most important piece of data. Including every single data point would make the graph very confusing.
1. The independent variable is type of treatment and the dependent variable is plant growth (in cm).
2. Type of treatment is not a continuous variable. There is no midpoint value between fertilizer brands (Brand A 1/2 doesn’t make sense). Plant growth is a continuous variable. It makes sense to sub-divide centimeters into smaller values. Since the independent variable is categorical and the dependent variable is continuous, this graph should be a bar graph.
3. Plant growth (the dependent variable) should go on the Y axis and type of treatment (the independent variable) should go on the X axis.
4. Notice that the values on the Y axis are continuous and evenly spaced. Each line represents an increase of 5cm.
5. Notice that both the X and the Y axis have labels that include units (when required).
6. Notice that the graph has a descriptive caption that allows the figure to stand alone without additional information given from the procedure: you know that this graph shows the average of the measurements taken from four tomato plants.
Descriptive captions
All figures that present data should stand alone – this means that you should be able to interpret the information contained in the figure without referring to anything else (such as the methods section of the paper). This means that all figures should have a descriptive caption that gives information about the independent and dependent variable. Another way to state this is that the caption should describe what you are testing and what you are measuring. A good starting point to developing a caption is “the effect of [the independent variable] on the [dependent variable].”
Here are some examples of good caption for figures:
• The effect of exercise on heart rate
• Growth rates of E. coli at different temperatures
• The relationship between heat shock time and transformation efficiency
Here are a few less effective captions:
• Heart rate and exercise
• Graph of E. coli temperature growth
• Table for experiment 1
Query \(4\) | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/01%3A_The_Process_of_Science/1.03%3A_Presenting_Data_-_Graphs_and_Tables.txt |
Whether scientific research is basic science or applied science, scientists must share their findings for other researchers to expand and build upon their discoveries. Communication and collaboration within and between sub disciplines of science are key to the advancement of knowledge in science. For this reason, an important aspect of a scientist’s work is disseminating results and communicating with peers. Scientists can share results by presenting them at a scientific meeting or conference, but this approach can reach only the limited few who are present. Instead, most scientists present their results in peer-reviewed articles that are published in scientific journals. Peer-reviewed articles are scientific papers that are reviewed, usually anonymously by a scientist’s colleagues, or peers. These colleagues are qualified individuals, often experts in the same research area, who judge whether or not the scientist’s work is suitable for publication. The process of peer review helps to ensure that the research described in a scientific paper or grant proposal is original, significant, logical, and thorough. Grant proposals, which are requests for research funding, are also subject to peer review. Scientists publish their work so other scientists can reproduce their experiments under similar or different conditions to expand on the findings. The experimental results must be consistent with the findings of other scientists.
There are many journals and the popular press that do not use a peer-review system. A large number of online open-access journals, journals with articles available without cost, are now available many of which use rigorous peer-review systems, but some of which do not. Results of any studies published in these forums without peer review are not reliable and should not form the basis for other scientific work. In one exception, journals may allow a researcher to cite a personal communication from another researcher about unpublished results with the cited author’s permission.
Scientific articles are not literary works. Instead, they are meant to transmit information effectively and concisely. The need for clarity and brevity is especially important for other forms of science communication such as posters where the audience must be able to understand the significance of your research in just a few minutes, but the need is there for all forms of scientific communication.
There is an explicit format that scientific papers follow, with relatively small variations in style among journals. Papers are broken down into the following sections: title, abstract, introduction, methods, results, and discussion. Every section, except the title, should be labeled as such. Generally the section name is centered and underlined (or bold-faced) over the text. Although posters follow the same format as a paper, each section is abbreviated (once again, clarity is critical).
Title
The title should give the reader a concise, informative description of the content and scope of the paper.
Abstract
The abstract is a concise summary of the major findings of the study. It should be no longer than 9-10 sentences. It should summarize every subsequent section of the paper. It should state the purposes of the study, and then briefly summarize the methods, results, and conclusions of the study. The abstract should be able to stand-alone. Do not refer to any figures or tables, or cite any references. Because the abstract is a distillation of the paper, it is often written last. It is typically the hardest part of the paper to write.
Introduction
In many journals, the introduction is also unlabeled, and simply starts after the abstract.
The introduction gives the rationale for the research. It answers the question “Why should anyone be interested in this work?” It usually includes background information, including the work of others, and a description of your objectives. If you are studying a particular species, give both the scientific (Latin) name and the common name the first time you mention your study animal. The scientific name is always underlined or italicized, and the genus name is capitalized while the species name is not. Cite only references pertinent to your study. Direct quotations are rarely used in scientific writing; instead state the findings of others in your own words. Furthermore, footnotes are rarely used in a scientific paper. Instead cite the author by last name, and the year that the source was published.
Smith (1987) found that male mice prefer the odor of non-pregnant female mice to that of pregnant female mice. Male mice prefer the odor of non-pregnant female mice to that of pregnant female mice (Smith, 1987).
When two people
co-author a paper, both are cited:
For instance:
When more than two people co-author a paper, cite only the first author, and refer to the other authors with the Latin phrase, “et al.”, indicating “and others”:
Undergraduate students who came to lectures were more likely to receive a high grade on the exams (Thatcher, et al., 2000).
Harrett and Garrett (1999) found no differences between male and female elephants in their response to the tape of a female vocalization.
The full reference for each work must be given in the literature cited section at the end of the paper. For references, select work from the primary literature: that is, work that is published by the same people who did it. In general, citing an encyclopedia or textbook is not appropriate for a scientific paper.
When organizing your introduction, begin with a general description of the topic, and then become more specific. For example, in a study of the olfaction in the reproductive behavior of mice, the skeleton of the introduction might be:
For reproduction to be successful, animals must be able to correctly assess the reproductive condition of a potential partner. Many different signals have evolved in animals to facilitate such assessment.
Olfactory signals seem to be particularly important in mammals.
Mice are particularly suited for studying the role of olfaction in reproductive behavior. Odor cues are involved in several aspects of mouse reproductive behavior, including… The aim of this study was…
Each of these sentences would be a good topic sentence of a different paragraph in the introduction. References should be cited where appropriate.
In sum, an introduction should convey your overall purpose in conducting the experiment as well as your specific objectives.
Methods
This section is also often called Materials and Methods.
This section is a very concise summary of the subjects, equipment, and procedures used. This section should contain enough information so that someone else could replicate your work. It is NOT a list, but a narrative description. Because it is a narrative, it should not include a list of your materials. Rather, they should be described in the narrative as required. For example, you could say: “We measured 5mL of enzyme solution into a test tube and heated it on a hot plate until it boiled.” From this, it is obvious that you used some sort of tool to accurately measure 5mL of solution, as well as a test tube, and a hot plate.
Only include information that is relevant to your experiment: do not include information that any scientist should know to do or that won’t affect the results (label the tubes, clean up afterwards, make a graph). If you are following the methods of another paper or a lab manual, simply cite the source. Then, you can concentrate on describing any changes that you made to that procedure. A common mistake is to let results creep into this section.
Results
The results includes presentations of your data and the results of statistical analysis of your data. First, state the overall trend of the data. Did the majority of the data statistically support or contradict the null hypothesis?
Address each statistical test separately, often in separate paragraphs. For each type of data analyzed say whether your results are statistically significant, and in parentheses give the statistical test used, the value of the test statistic, and the probability level for that computed value. For example, “Male mice visited non-pregnant females significantly more often than pregnant females (chi square = 4.69; p < 0.05).”
Do not present your raw data. Instead, present data in an easy to read form. You will probably use a figure or a table to present your results. Refer to each table by a number (Table 1, Table 2, etc.) It should have a concise heading at the top. Graphs and diagrams are both called figures and are numbered consecutively (Fig. 1, Fig. 2, etc.) They have headings at the bottom. Axes on graphs should be clearly labeled. See the section on Presenting Your Data for more information.
You must refer to every table and figure at least once in the text. Often this can be done parenthetically: “Male mice visited non-pregnant females significantly more often than pregnant females (chi square = 4.69; p <0.05; Fig. 2).”
Do not use the word “significant” unless it can be supported by statistical evidence.
A common mistake is to discuss the implications of your findings. Save that for the discussion section.
Discussion
Here you are to give a reader the “take home” message of the study. Begin by briefly summarizing the major findings of your study. Then discuss each finding one at a time (usually in separate paragraphs).
Interpret your results in light of the biology you are studying. Your discussion section should parallel your introduction: if you discussed the role of reproductive biology of the mouse at the beginning of your study, come back to it again here. The paper should come full circle.
Use references throughout your discussion to support your points. Compare your findings with those of similar studies.
Do not make statements that cannot be supported by the data, and be sure none of your conclusions are contradicted by the data. Discuss unexpected results or possible errors in the experiment, but don’t focus on “what didn’t work”. We all know this was a classroom research project!
Literature cited
Each academic discipline uses a different format to cite the references they use. These differences can be dramatic (English vs. Science, for example) or small (Psychology vs. Biology), but they are based on what information is seen as important. In this course, we follow the format of the most biology journals by using CSE format. See the section on Citing Your Sources for more specific information.
General hints
For stylistic hints, browse one of the many books in the library on scientific writing. Remember, being a good writer in English “121” doesn’t mean your skills will translate to science writing without work (though you have a great start!).
Outline your paper. Use topic sentences for every paragraph. You should be able to go back and underline each topic sentence after you are finished.
Keep your report as short as you can, consistent with clarity and completeness. Do not “pad” with a lot of irrelevant information just to show you know a lot.
A note on Plagiarism: Plagiarism is a serious academic offense. However, most instances of plagiarism are the result of a lack of care and effort, and not intentional misbehavior. Here is a general rule to follow: Don’t Cut and Paste! Accidental or not, any occurrence of academic dishonesty will be treated seriously. Ignorance is no defense.
Be sure to proofread for typographical errors, poor grammar, or unclear sentence structure.
Try to start paragraphs with a topic sentence or a summary statement. Then follow it with supporting statements. This technique makes your writing clearer and easier to follow. Ideally, someone could read the first sentence of each paragraph and still understand the gist of our paper.
PLEASE avoid dull scientific writing, particularly the use of the passive voice. As much as possible, use an active voice. Passive writing takes up more space and is dull, dull, dull. Look at the example here; see how this is more exciting and can lead to an interesting ecological observation about the importance of the predator – prey relationship involved?
BAD: Mussels are eaten by sea stars.
GOOD: Sea stars eat mussels.
BETTER STILL: Sea stars are voracious predators of mussels.
Make sure the object to which words such as “this” or “it” refer is clear.
Combine sentences with low information content into one sentence. This will make your writing more streamlined and less repetitive. But don’t write run-on sentences either!
Always refer to work people have done in the past in the past tense. Refer to species attributes or other ongoing, continuing states in the present tense.
The word “data” is plural. Say either “these data are…” or “this datum is…”
BAD: Wentworth (1985) studied vegetation in Arizona. He found that tree species distributions followed gradients.
GOOD: In the Huachuca Mountains of Arizona, both elevation and the amount of light influenced tree species distributions (Wentworth 1985).
Scientific names of animals and plants are underlined or italicized (as are most Latin words), such as Homo sapiens or Homo sapiens (genera and all higher taxa are capitalized, species names are lowercase).
Do not anthropomorphize. A honeybee or a dandelion does not have the same consciousness or emotional life as your roommate. In extreme forms, this type of writing is appropriate for the tabloids in supermarket checkout lines…
Try varying the length of your sentences, and keep in mind that a sentence with 4 words is probably too short, and one with 20 too long.
BAD: Kudzu, an Asian super weed, intends to dominate and conquer the entire southeastern United States.
GOOD: Kudzu is a noxious weed introduced from Asia that has quickly spread from its point of introduction throughout the southeastern United States.
Avoid using too many clauses in one sentence. If you see that you
have a lot of commas, that is a clue that you’ve overdone the number of clauses in the sentence.
Try reading your work out loud. Anything that is written poorly will be difficult to read. This technique will alert you to problem areas in your writing.
BE PREPARED TO WRITE SEVERAL DRAFTS! Good, hard editing will turn you from a mediocre to a good writer. And with good writing, you are able to show your GREAT thinking!
Sources:
These instructions are adapted by Walter Shriner. Originally from Jakob, E. 1995. Laboratory manual for animal behavior. Bowling Green University and Muller, K. 1991. Ornithology laboratory. University of California, Davis.
Unless otherwise noted, images on this page are licensed under CC-BY 4.0 by OpenStax.
OpenStax, Biology. OpenStax CNX. May 27, 2016 http://cnx.org/contents/[email protected]:RD6ERYiU@5/The-Process-of-Science. | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/01%3A_The_Process_of_Science/1.04%3A_Writing_for_Science.txt |
Learning Objectives
Course Objective for this section: Select, evaluate, and utilize discipline-specific information and literature to explore topics.
• Differentiate between questions that can and cannot be answered using science.
• Identify appropriate credible sources of information to research a topic.
• Evaluate sources of information for their strengths and weaknesses.
Science is a very specific way of learning, or knowing, about the world. Humans have used the process of science to learn a huge amount about the way the natural world works. Science is responsible for amazing innovations in medicine, hygiene, and technology. There are however, areas of knowledge and human experience that the methods of science cannot be applied to. These include such things as answering purely moral questions, aesthetic questions, or what can be generally categorized as spiritual questions. Science has cannot investigate these areas because they are outside the realm of material phenomena, the phenomena of matter and energy, and cannot be observed and measured.
Questions that can be answered using science Questions that cannot be answered using science
• What is the optimum temperature for the growth of E. coli bacteria?
• Do birds prefer bird feeders of a specific color?
• What is the cause of this disease?
• How effective is this drug in treating this disease?
• How tall is Santa Claus?
• Do angels exist?
• Which is better: classical music or rock and roll?
• What are the ethical implications of human cloning?
Since this is a biology class, we will be focusing on questions that can be answered scientifically. Remember that in the scientific process, observations lead to questions. A scientific question is one that can be answered by using the process of science (testing hypotheses, making observations about the natural world, designing experiments).
Sometimes you will directly make observations yourself about the natural world that lead you to ask scientific questions, other times you might hear or read something that leads you to ask a question. Regardless of how you make your initial observation, you will want to do research about your topic before you start setting up an experiment. When you’re learning about a topic, it’s important to use credible sources of information.
Types of Sources
Whether conducting research in the social sciences, humanities (especially history), arts, or natural sciences, the ability to distinguish between primary and secondary source material is essential. Basically, this distinction illustrates the degree to which the author of a piece is removed from the actual event being described. This means whether the author is reporting information first hand (or is first to record these immediately following an event), or conveying the experiences and opinions of others—that is, second hand. In biology, the distinction would be between the person (or people) who conducted the research and someone who didn’t actually do the research, but is merely reporting on it.
Primary sources
These are contemporary accounts of an event, written by someone who experienced or witnessed the event in question. In general, these original documents (i.e., they are not about another document or account) are often diaries, letters, memoirs, journals, speeches, manuscripts, interviews, photographs, audio or video recordings, or original literary or theatrical works.
In science, a “primary source” or the “primary literature” refers to the original publication of a scientist’s new data, results, and conclusions. These articles are written for other experts in a specific scientific field.
You’ve probably done a writing assignment or other project during which you have participated in a peer review process. During this process, your project was critiqued and evaluated by people of similar competence to yourself (your peers). This gave you feedback on which to improve your work. Scientific articles typically go through a peer review process before they are published in an academic journal. In this case, the peers who are reviewing the article are other experts in the specific field about which the paper is written. This allows other scientists to critique experimental design, data, and conclusions before that information is published in an academic journal. Often, the scientists who did the experiment and who are trying to publish it are required to do additional work or edit their paper before it is published. The goal of the scientific peer review process is to ensure that published primary articles contain the best possible science.
Secondary sources
The function of a secondary source is to interpret the primary source. A secondary source can be described as at least one step removed from the event or phenomenon under review. Secondary source materials interpret, assign value to, conjecture upon, and draw conclusions about the events reported in primary sources. These are usually in the form of published works such as magazine articles or books, but may include radio or television documentaries, or conference proceedings.
Popular vs. Scholarly Sources
POPULAR SCHOLARLY
Broad range of topics, presented in shorter articles Specific, narrowly focused topics in lengthy, in-depth articles
Articles offer overview of subject matter; interpretation, rather than original research; sometimes contain feature articles and reports on current social issues and public opinion Articles often contain previously unpublished research and detail new developments in field
Intended to attract a general readership without any particular expertise or advanced education Intended for specialist readership of researchers, academics, students and professionals
Written by staff (not always attributed) or freelance writers using general, popular language Written by identified specialists and researchers in subject area, usually employing technical, subject-specific language and jargon
Edited and approved for publication in-house (not peer-reviewed) Critically evaluated by peers (fellow scholars) in field for content, scholarly soundness, and academic value
Articles rarely contain references or footnotes and follow no specific format Well-researched, documented articles nearly always follow standard format:
abstract, introduction, literature review, methodology, results, conclusion, bibliography/references
Designed to attract eye of potential newsstand customers: usually filled with photographs or illustrations, printed on glossier paper Sober design: mostly text with some tables or graphs accompanying articles; usually little or no photography; negligible, if any, advertising; rarely printed on high-gloss paper
Each issue begins with page number ‘1’ Page numbers of issues within a volume (year) are usually consecutive (i.e., first page of succeeding issue is number following last page number of previous issue)
Presented to entertain, promote point of view, and/or sell products Intended to present researchers’ opinions and findings based on original research
Examples: Newsweek, Rolling Stone, Vogue Examples: Science, Nature, Journal of Microbial and Biochemical Technology
In science, it is often extremely difficult to read and understand primary articles unless you are an expert in that specific scientific field. Secondary sources are typically easier to read and can give you the important information from a primary source, but only if the secondary source has interpreted the information correctly! It is always better to go to the primary source if possible because otherwise you are relying on someone else’s interpretation of the information. However, it is always better to use a source that you can read and understand rather than a source that you can’t. For this reason, it is very important to be able to identify credible secondary sources.
Query \(1\)
Evaluating Credibility
When you write a scientific paper (or any paper, really), you want to back up your statements with credible sources. You will need to identify credible sources to help you research scientific topics to help you develop interesting scientific questions. You will also need sources to help you form a well-educated hypothesis that is not just based on your guess about what will happen. A credible source is one that is trustworthy from which the information can be believed. Credible sources are written by people who are experts in the field (or at least are very knowledgeable) about the subject that they are commenting on.
We will be using a variation of the CRAAP test to help you determine whether or not sources that you find are credible or not. The CRAAP Test was created by Sarah Blakeslee, of the University of California at Chico’s Meriam Library. It is adapted below. When evaluating the credibility of sources using this method, if it’s CRAAP, it’s good!
You can use the table below to help you evaluate the credibility of your sources.
Credibility Table
Factors to consider Least reliable
(0 points)
Possibly reliable
(1 point)
Most reliable
(2 points)
Currency No date of publication or revision given Outdated for this particular topic Recently published or revised
Reliable source Unreliable website, no additional info available Possibly reliable Official government or organization, institutional sites, academic journals
Author No author is given / the author is not qualified to write about this topic Author is educated on topic or is staff of an organization assumed to be knowledgeable on this specific topic Specifically identified expert in this field with degrees / credentials in this subject
Accuracy No review process and information is not supported by evidence from cited sources The information may have been reviewed or edited by someone knowledgeable in the field. It mentions but does not directly cite other sources The information has been peer reviewed and is supported by evidence from cited credible sources
Purpose Obviously biased or trying to sell you something Sponsored source; may present unbalanced information Balanced, neutral, presents all sides of the issue fully
In general, do not use a source if it doesn’t pass the CRAAP test! For our purposes, do not use any sources that score less than 6 points using the credibility table.
Several examples are given below for sources that you might come across if you were researching the topic of vaccine safety.
Example 1:
CDC (Centers for Disease Control and Prevention). Aug 28, 2015. Vaccine Safety [Internet]. [cited May 12, 2016]. Available from: http://www.cdc.gov/vaccinesafety/index.html
Score Discussion – why did you give that score?
Currency 2 Aug 28 2015 is recent and shows that this information is updated frequently.
Reliable source 2 I looked at the “about this organization” and learned that the CDC is a major government organization that works to protect Americans from health, safety, and security threats. They are a division of the US department of health and human services.
Author 1 A specific author was not identified, but the page states that the content is from the CDC, which suggests that it was written by a knowledgeable staff member.
Accuracy 1.5 No information is given about the review process, but it was probably edited by staff at the CDC. There is a list of citations and links to primary scientific articles supporting the information.
Purpose 2 The point of view does not appear to be biased because it seems to be presenting factual information. Admittedly, it only presents the pro-vaccine side of the argument. There are no ads on the page or other information trying to change the reader’s viewpoint.
Credibility Score 8.5/10 This seems like an excellent source to use for research. It’s readable and I could look at the primary articles if I wanted to check them out.
Example 2:
Stop Mandatory Vaccination. N.d.. The Dangers of Vaccines and Vaccinations [Internet]. [cited May 12, 2016]. Available from: http://www.stopmandatoryvaccination.com/vaccine-dangers/
Score Discussion – why did you give that score?
Currency 1 The copyright is given as 2015, but there is no date for this specific article. It does reference something that took place in 2015, so it is likely written after that.
Reliable source 0 The “About” page states that the organization was started by Larry Cook using a GoFundMe platform
Author 0 Larry Cook has been devoted to the natural lifestyle for 25 years, but doesn’t appear to have any degrees or specific expertise on this topic. Other contributing authors include Landee Martin, who has a Bachelor’s of Science in Psychology (which isn’t related to vaccine safety), and Brittney Kara, who is a mother who has studied holistic living for the last 17 years. None of the individuals specifically identified on the website appear to be experts in the field.
Accuracy 0.5 It seems unlikely that there is any review process. There are links to several sources, but none of them appear to be primary scientific articles. Several are links to interviews.
Purpose 0 This source is extremely biased. Even the name of the website is biased. There is a link to donate to the webpage. There are at least 10 ads for anti-vaccine books and websites.
Credibility Score 1.5/10 I would not want to use this source to research this topic. It’s extremely biased and doesn’t seem to offer much evidence for its assertions.
Citing Your Sources
One of the goals for any class is to help students become better scholars. And, one of the important skills of scholarship is proper citation of resources used. Citations demonstrate your “credentials” as a scholar, and provide a resource to your readers of good reference material.
Why do you have to cite your sources?
No research paper is complete without a list of the sources that you used in your writing. Scholars are very careful to keep accurate records of the resources they’ve used, and of the ideas and concepts they’ve quoted or used from others. This record keeping is generally presented in the form of citations.
A citation is a description of a book, article, URL, etc. that provides enough information so that others can locate the source you used themselves. It allows you to credit the authors of the sources you use and clarify which ideas belong to you and which belong to other sources. And providing a citation or reference will allow others to find and use these sources as well. Most research papers have a list of citations or cited references and there are special formatting guidelines for different types of research.
However, there are many “proper” formats because each discipline has its own rules. In general we ask only that you use one of the “official” formats and that you use it consistently. To understand what we mean by “consistent”, compare the citations in two scientific journals. You will notice that each journal has its own rules for whether an article title is in quotes, bold, underlined, etc., but within each journal the rule applies to all reference citations. Below is a condensed guide to the general format used in science (CSE). For more detailed information consult one of the online citation guides and generators.
Plagiarism
Plagiarism is presenting the words or ideas of someone else as your own without proper acknowledgment of the source. When you work on a research paper you will probably find supporting material for your paper from works by others. It’s okay to quote people and use their ideas, but you do need to correctly credit them. Even when you summarize or paraphrase information found in books, articles, or Web pages, you must acknowledge the original author. To avoid plagiarism, include a reference to any material you use that provides a fact not commonly known, or whenever you use information from another author. In short, if you didn’t collect the data or reach the conclusion on your own, cite it!
These are all examples of plagiarism:
• Buying or using a term paper written by someone else.
• Cutting and pasting passages from the Web, a book, or an article and inserting them into your paper without citing them. Warning! It is now easy for your instructors to search and identify passages that you have copied from the Web.
• Using the words or ideas of another person without citing them.
• Paraphrasing that person’s words without citing them.
Tips for Avoiding Plagiarism:
• First, use your own ideas—it should be your paper and your ideas should be the focus.
• Use the ideas of others sparingly—only to support or reinforce your own argument.
• When taking notes, include complete citation information for each item you use.
• Use quotation marks when directly stating another person’s words. Quotes are not frequently used in scientific writing unless you are directly quoting someone’s spoken words.
Citing Sources in CSE Format
The Council of Science Editors (CSE) citation format is commonly used in scientific writing. CSE format emphasizes the information that is important when writing scientifically: who wrote the information and when they wrote it. In different fields, there is an emphasis on different types of information. In the humanities, MLA format is commonly used. This style emphasizes the author’s name and the page number. This information allows a reader to track down the exact quotes that are being discussed. Another commonly used format, APA, emphasizes the author’s name and the year the information was published.
The standard format for citing a source in science writing is the Name-year format. In this format, the first author’s last name is followed by the date. For example: Not all populations of alligators in the everglades are at risk from habitat loss (Nicholson, 2002).
If you are not familiar with the CSE citation style, you can get additional information and examples at http://writing.wisc.edu/Handbook/DocCSE_NameYear.html
Beware of computerized “citation creators.” While they can get you part way to a correct citation, they rarely are 100% correct. For example, they often fail to put the last name first.
Citing a scientific journal article
Author’s last name first initial, next author’s last name first initial. Date published. Title of Article. Journal Name. Volume (issue): pages.
Please note that you need to cite the JOURNAL, not the DATABASE that you got it from. Citing the database in which you found a scientific journal article is like citing Google for an internet resource that you are using.
Flores-Cruz Z, Allen C. 2011. Necessity of OxyR for the hydrogen peroxide stress response and full virulence in Ralstonia solanacearum. Appl Environ Microbiol. 77(18):6426-6432.
Werling BP, Lowenstein DM, Straub CS, Gratton C. 2012. Multi-predator effects produced by functionally distinct species vary with prey density. J Insect Sci; 12(30): 346-378.
Shriner, W.M. 1998. Yellow-bellied marmot and golden-mantled ground squirrel responses to heterospecific alarm calls. Animal Behaviour 55:529-536.
Citing an internet resource
Author’s last name, first initial. Date published. Title of Website [Internet]. Publisher information. [cited on date that you accessed the information]. Available from: URL where you accessed the source.
Williamson RC. 2004. Deciduous tree galls [Internet]. Madison (WI): University of Wisconsin-Madison; [cited 2013 Sep 12]. Available from: http://labs.russell.wisc.edu/pddc/files/Fact_Sheets/FC_PDF/Deciduous_Tree_Galls.pdf
[BP] The Biology Project. 2003. The chemistry of amino acids [Internet]. University of Arizona; [cited 2004 Mar 17]. Available from: http://www.biology.arizona.edu/biochemistry/problem_sets/aa/aa.html
Hilton-Taylor C, compiler. 2000. 2000 IUCN red list of threatened species [Internet]. Gland (Switzerland) and Cambridge (UK): IUCN; [cited 2002 Feb 12]. Available from: http://www.redlist.org/.
Query \(2\)
Query \(3\)
Citing Sources Within Text
We will be using the CSE Name-year format for citations. When you want to provide a citation reference for a statement that you are making, you should end the sentence with (First author’s last name, year). If the article was written by an organization and not a specific author, you can use the name of the organization (or an abbreviation for the name).
Example: Sickle cell anemia is caused by abnormally-shaped haemoglobin proteins (NIH, 2012).
In the References Cited section (a.k.a. Literature cited…) list all the sources you cited in your paper, but do not include any items that you did not specifically cite within the body of your paper or project, even if you read them! Except in rare instances, do not cite a reference that you have not personally read.
You should then list all your references in the Literature Cited section alphabetically by author’s last name.
For more information and lots of examples of what to do in specific instances, please visit http://writing.wisc.edu/Handbook/DocCSE_NameYear.html | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/01%3A_The_Process_of_Science/1.05%3A_Using_Credible_Sources.txt |
Learning Outcomes
Course Outcomes for this section:
• Describe the structure of biologically-important molecules (carbohydrates, lipids, proteins, nucleic acids, water) and how their structure leads to their function.
Living things are highly organized and structured, following a hierarchy that can be examined on a scale from small to large. The examination of the smallest parts involves a knowledge of chemistry. We can put the levels of organization of living things in order from smallest to largest.
• The atom is the smallest and most fundamental unit of matter. It consists of a nucleus surrounded by electrons.
• Atoms form molecules. A molecule is a chemical structure consisting of at least two atoms held together by one or more chemical bonds. Many molecules that are biologically important are macromolecules, large molecules that are typically formed by polymerization (a polymer is a large molecule that is made by combining smaller units called monomers, which are simpler than macromolecules).
These first 2 levels (or 3, depending on how you categorize macromolecules) are typically studied in chemistry or biochemistry courses. However, a working knowledge of atoms and molecules is required to understand how these small pieces work to make larger, living organisms.
• Some cells contain aggregates of macromolecules surrounded by membranes; these are called organelles. Organelles are small structures that exist within cells.
• All living things are made of cells; the cell itself is the smallest fundamental unit of structure and function in living organisms.
• In larger multicellular organisms, cells combine to make tissues, which are groups of similar cells carrying out similar or related functions.
• Organs are collections of tissues grouped together performing a common function. An organ system is a higher level of organization that consists of functionally related organs.
• Mammals have many organ systems. For instance, the circulatory system transports blood through the body and to and from the lungs; it includes organs such as the heart and blood vessels.
• Organisms are individual living entities. For example, each tree in a forest is an organism. Single-celled prokaryotes and single-celled eukaryotes are also considered organisms and are typically referred to as microorganisms.
Once we move beyond one single organism, we have reached the study of ecology. You’ll look at these topics in BI213.
• All the individuals of a species living within a specific area are collectively called a population. For example, a forest may include many pine trees. All of these pine trees represent the population of pine trees in this forest. Different populations may live in the same specific area. For example, the forest with the pine trees includes populations of flowering plants and also insects and microbial populations.
• A community is the sum of populations inhabiting a particular area. For instance, all of the trees, flowers, insects, and other populations in a forest form the forest’s community. The forest itself is an ecosystem.
• An ecosystem consists of all the living things in a particular area together with the abiotic, non-living parts of that environment such as nitrogen in the soil or rain water.
• At the highest level of organization, the biosphere is the collection of all ecosystems, and it represents the zones of life on earth. It includes land, water, and even the atmosphere to a certain extent.
02: Chemistry for Biology
An atom is the smallest component of an element that retains all of the chemical properties of that element. For example, one hydrogen atom has all of the properties of the element hydrogen, such as it exists as a gas at room temperature, and it bonds with oxygen to create a water molecule. Hydrogen atoms cannot be broken down into anything smaller while still retaining the properties of hydrogen. If a hydrogen atom were broken down into subatomic particles, it would no longer have the properties of hydrogen.
All atoms contain protons, electrons, and neutrons (Figure \(1\)). The only exception is hydrogen (H), which is made of one proton and one electron. A proton is a positively charged particle that resides in the nucleus (the core of the atom) of an atom and has a mass of 1 and a charge of +1. An electron is a negatively charged particle that travels in the space around the nucleus. In other words, it resides outside of the nucleus. It has a negligible mass and has a charge of –1.
Neutrons, like protons, reside in the nucleus of an atom. They have a mass of 1 and no charge. The positive (protons) and negative (electrons) charges balance each other in a neutral atom, which has a net zero charge.
Because protons and neutrons each have a mass of 1, the mass of an atom is equal to the number of protons and neutrons of that atom. The number of electrons does not factor into the overall mass, because their mass is so small.
Query \(1\)
At the most basic level, all organisms are made of a combination of elements. An element is a substance whose atoms all have the same number of protons. They contain atoms that combine together to form molecules. In multicellular organisms, such as animals, molecules can interact to form cells that combine to form tissues, which make up organs. These combinations continue until entire multicellular organisms are formed.
Each element has its own unique properties. Each contains a different number of protons and neutrons, giving it its own atomic number and mass number. The atomic number of an element is equal to the number of protons that element contains. The mass number, or atomic mass, is the number of protons plus the number of neutrons of that element. Therefore, it is possible to determine the number of neutrons by subtracting the atomic number from the mass number.
These numbers provide information about the elements and how they will react when combined. Different elements have different melting and boiling points, and are in different states (liquid, solid, or gas) at room temperature. They also combine in different ways. Some form specific types of bonds, whereas others do not. How they combine is based on the number of electrons present. Because of these characteristics, the elements are arranged into the periodic table of elements, a chart of the elements that includes the atomic number and relative atomic mass of each element. The periodic table also provides key information about the properties of elements (Figure \(2\)) —often indicated by color-coding. The arrangement of the table also shows how the electrons in each element are organized and provides important details about how atoms will react with each other to form molecules.
Isotopes are different forms of the same element that have the same number of protons, but a different number of neutrons. Some elements, such as carbon, potassium, and uranium, have naturally occurring isotopes. Carbon-12, the most common isotope of carbon, contains six protons and six neutrons. Therefore, it has a mass number of 12 (six protons and six neutrons) and an atomic number of 6 (which makes it carbon). Carbon-14 contains six protons and eight neutrons. Therefore, it has a mass number of 14 (six protons and eight neutrons) and an atomic number of 6, meaning it is still the element carbon. These two alternate forms of carbon are isotopes. Some isotopes are unstable and will lose protons, other subatomic particles, or energy to form more stable elements. These are called radioactive isotopes or radioisotopes.
Query \(2\)
Evolution in Action: Carbon Dating
Carbon-14 (14C) is a naturally occurring radioisotope that is created in the atmosphere by cosmic rays. This is a continuous process, so more 14C is always being created. As a living organism develops, the relative level of 14C in its body is equal to the concentration of 14C in the atmosphere. When an organism dies, it is no longer ingesting 14C, so the ratio will decline. 14C decays to 14N by a process called beta decay; it gives off energy in this slow process.
After approximately 5,730 years, only one-half of the starting concentration of 14C will have been converted to 14N. The time it takes for half of the original concentration of an isotope to decay to its more stable form is called its half-life. Because the half-life of 14C is long, it is used to age formerly living objects, such as fossils. Using the ratio of the 14C concentration found in an object to the amount of 14C detected in the atmosphere, the amount of the isotope that has not yet decayed can be determined. Based on this amount, the age of the fossil can be calculated to about 50,000 years (Figure \(3\)). Isotopes with longer half-lives, such as potassium-40, are used to calculate the ages of older fossils. Through the use of carbon dating, scientists can reconstruct the ecology and biogeography of organisms living within the past 50,000 years.
Query \(3\) | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/02%3A_Chemistry_for_Biology/2.01%3A_Atoms.txt |
How elements interact with one another depends on how their electrons are arranged and how many openings for electrons exist at the outermost region where electrons are present in an atom. Electrons exist at energy levels that form shells around the nucleus. The closest shell can hold up to two electrons. The closest shell to the nucleus is always filled first, before any other shell can be filled. Hydrogen has one electron; therefore, it has only one spot occupied within the lowest shell. Helium has two electrons; therefore, it can completely fill the lowest shell with its two electrons. If you look at the periodic table, you will see that hydrogen and helium are the only two elements in the first row. This is because they only have electrons in their first shell. Hydrogen and helium are the only two elements that have the lowest shell and no other shells.
The second and third energy levels can hold up to eight electrons. The eight electrons are arranged in four pairs and one position in each pair is filled with an electron before any pairs are completed.
Looking at the periodic table again (Figure 2.1.2), you will notice that there are seven rows. These rows correspond to the number of shells that the elements within that row have. The elements within a particular row have increasing numbers of electrons as the columns proceed from left to right. Although each element has the same number of shells, not all of the shells are completely filled with electrons. If you look at the second row of the periodic table, you will find lithium (Li), beryllium (Be), boron (B), carbon (C), nitrogen (N), oxygen (O), fluorine (F), and neon (Ne). These all have electrons that occupy only the first and second shells. Lithium has only one electron in its outermost shell, beryllium has two electrons, boron has three, and so on, until the entire shell is filled with eight electrons, as is the case with neon.
Not all elements have enough electrons to fill their outermost shells, but an atom is at its most stable when all of the electron positions in the outermost shell are filled. Because of these vacancies in the outermost shells, we see the formation of chemical bonds, or interactions between two or more of the same or different elements that result in the formation of molecules. To achieve greater stability, atoms will tend to completely fill their outer shells and will bond with other elements to accomplish this goal by sharing electrons, accepting electrons from another atom, or donating electrons to another atom. Because the outermost shells of the elements with low atomic numbers (up to calcium, with atomic number 20) can hold eight electrons, this is referred to as the octet rule. An element can donate, accept, or share electrons with other elements to fill its outer shell and satisfy the octet rule.
When an atom does not contain equal numbers of protons and electrons, it is called an ion. Because the number of electrons does not equal the number of protons, each ion has a net charge. Positive ions are formed by losing electrons and are called cations. Negative ions are formed by gaining electrons and are called anions.
For example, sodium only has one electron in its outermost shell. It takes less energy for sodium to donate that one electron than it does to accept seven more electrons to fill the outer shell. If sodium loses an electron, it now has 11 protons and only 10 electrons, leaving it with an overall charge of +1. It is now called a sodium ion.
The chlorine atom has seven electrons in its outer shell. Again, it is more energy-efficient for chlorine to gain one electron than to lose seven. Therefore, it tends to gain an electron to create an ion with 17 protons and 18 electrons, giving it a net negative (–1) charge. It is now called a chloride ion. This movement of electrons from one element to another is referred to as electron transfer. As Figure \(1\) illustrates, a sodium atom (Na) only has one electron in its outermost shell, whereas a chlorine atom (Cl) has seven electrons in its outermost shell. A sodium atom will donate its one electron to empty its shell, and a chlorine atom will accept that electron to fill its shell, becoming chloride. Both ions now satisfy the octet rule and have complete outermost shells. Because the number of electrons is no longer equal to the number of protons, each is now an ion and has a +1 (sodium) or –1 (chloride) charge.
Ionic Bonds
There are four types of bonds or interactions: ionic, covalent, hydrogen bonds, and van der Waals interactions. Ionic and covalent bonds are strong interactions that require a larger energy input to break apart. When an element donates an electron from its outer shell, as in the sodium atom example above, a positive ion is formed (Figure \(2\)). The element accepting the electron is now negatively charged. Because positive and negative charges attract, these ions stay together and form an ionic bond, or a bond between ions. The elements bond together with the electron from one element staying predominantly with the other element. When Na+ and Cl ions combine to produce NaCl, an electron from a sodium atom stays with the other seven from the chlorine atom, and the sodium and chloride ions attract each other in a lattice of ions with a net zero charge.
Covalent Bonds
Another type of strong chemical bond between two or more atoms is a covalent bond. These bonds form when an electron is shared between two elements and are the strongest and most common form of chemical bond in living organisms. Covalent bonds form between the elements that make up the biological molecules in our cells. Unlike ionic bonds, covalent bonds do not dissociate in water.
Interestingly, chemists and biologists measure bond strength in different ways. Chemists measure the absolute strength of a bond (the theoretical strength) while biologists are more interested in how the bond behaves in a biological system, which is usually aqueous (water-based). In water, ionic bonds come apart much more readily than covalent bonds, so biologists would say that they are weaker than covalent bonds. If you look in a chemistry textbook, you’ll see something different. This is a great example of how the same information can lead to different answers depending on the perspective that you’re viewing it from.
The hydrogen and oxygen atoms that combine to form water molecules are bound together by covalent bonds. The electron from the hydrogen atom divides its time between the outer shell of the hydrogen atom and the incomplete outer shell of the oxygen atom. To completely fill the outer shell of an oxygen atom, two electrons from two hydrogen atoms are needed, hence the subscript “2” in H2O. The electrons are shared between the atoms, dividing their time between them to “fill” the outer shell of each. This sharing is a lower energy state for all of the atoms involved than if they existed without their outer shells filled.
There are two types of covalent bonds: polar and nonpolar. Nonpolar covalent bonds form between two atoms of the same element or between different elements that share the electrons equally. For example, an oxygen atom can bond with another oxygen atom to fill their outer shells. This association is nonpolar because the electrons will be equally distributed between each oxygen atom. Two covalent bonds form between the two oxygen atoms because oxygen requires two shared electrons to fill its outermost shell. Nitrogen atoms will form three covalent bonds (also called triple covalent) between two atoms of nitrogen because each nitrogen atom needs three electrons to fill its outermost shell. Another example of a nonpolar covalent bond is found in the methane (CH4) molecule. The carbon atom has four electrons in its outermost shell and needs four more to fill it. It gets these four from four hydrogen atoms, each atom providing one. These elements all share the electrons equally, creating four nonpolar covalent bonds (Figure \(3\)).
In a polar covalent bond, the electrons shared by the atoms spend more time closer to one nucleus than to the other nucleus. Because of the unequal distribution of electrons between the different nuclei, a slightly positive (δ+) or slightly negative (δ–) charge develops. The covalent bonds between hydrogen and oxygen atoms in water are polar covalent bonds. The shared electrons spend more time near the oxygen nucleus, giving it a small negative charge, than they spend near the hydrogen nuclei, giving these molecules a small positive charge.
Hydrogen Bonds
Ionic and covalent bonds are strong bonds that require considerable energy to break. However, not all bonds between elements are ionic or covalent bonds. Weaker bonds can also form. These are attractions that occur between positive and negative charges that do not require much energy to break. Two weak bonds that occur frequently are hydrogen bonds and van der Waals interactions. These bonds give rise to the unique properties of water and the unique structures of DNA and proteins.
When polar covalent bonds containing a hydrogen atom form, the hydrogen atom in that bond has a slightly positive charge. This is because the shared electron is pulled more strongly toward the other element and away from the hydrogen nucleus. Because the hydrogen atom is slightly positive (δ+), it will be attracted to neighboring negative partial charges (δ–). When this happens, a weak interaction occurs between the δ+ charge of the hydrogen atom of one molecule and the δ– charge of the other molecule. This interaction is called a hydrogen bond. This type of bond is common; for example, the liquid nature of water is caused by the hydrogen bonds between water molecules (Figure \(4\)). Hydrogen bonds give water the unique properties that sustain life. If it were not for hydrogen bonding, water would be a gas rather than a liquid at room temperature.
Hydrogen bonds can form between different molecules and they do not always have to include a water molecule. Hydrogen atoms in polar bonds within any molecule can form bonds with other adjacent molecules. For example, hydrogen bonds hold together two long strands of DNA to give the DNA molecule its characteristic double-stranded structure. Hydrogen bonds are also responsible for some of the three-dimensional structure of proteins.
van der Waals Interactions
Like hydrogen bonds, van der Waals interactions are weak attractions or interactions between molecules. They occur between polar, covalently bound, atoms in different molecules. Some of these weak attractions are caused by temporary partial charges formed when electrons move around a nucleus. These weak interactions between molecules are important in biological systems.
Query \(1\)
Query \(2\) | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/02%3A_Chemistry_for_Biology/2.02%3A_Chemical_Bonds.txt |
Do you ever wonder why scientists spend time looking for water on other planets? It is because water is essential to life; even minute traces of it on another planet can indicate that life could or did exist on that planet. Water is one of the more abundant molecules in living cells and the one most critical to life as we know it. Approximately 60–70 percent of your body is made up of water. Without it, life simply would not exist.
Water Is Polar
The hydrogen and oxygen atoms within water molecules form polar covalent bonds. The shared electrons spend more time associated with the oxygen atom than they do with hydrogen atoms. There is no overall charge to a water molecule, but there is a slight positive charge on each hydrogen atom and a slight negative charge on the oxygen atom. Because of these charges, the slightly positive hydrogen atoms repel each other and form the unique shape. Each water molecule attracts other water molecules because of the positive and negative charges in the different parts of the molecule.
Water also attracts other polar molecules (such as sugars), forming hydrogen bonds. When a substance readily forms hydrogen bonds with water, it can dissolve in water and is referred to as hydrophilic (“water-loving”). Hydrogen bonds are not readily formed with nonpolar substances like oils and fats (Figure \(3\)). These nonpolar compounds are hydrophobic (“water-fearing”) and will not dissolve in water.
Water Stabilizes Temperature
The hydrogen bonds in water allow it to absorb and release heat energy more slowly than many other substances. Temperature is a measure of the motion (kinetic energy) of molecules. As the motion increases, energy is higher and thus temperature is higher. Water absorbs a great deal of energy before its temperature rises. Increased energy disrupts the hydrogen bonds between water molecules. Because these bonds can be created and disrupted rapidly, water absorbs an increase in energy and temperature changes only minimally. This means that water moderates temperature changes within organisms and in their environments. As energy input continues, the balance between hydrogen-bond formation and destruction swings toward the destruction side. More bonds are broken than are formed. This process results in the release of individual water molecules at the surface of the liquid (such as a body of water, the leaves of a plant, or the skin of an organism) in a process called evaporation. Evaporation of sweat, which is 90 percent water, allows for cooling of an organism, because breaking hydrogen bonds requires an input of energy and takes heat away from the body.
Conversely, as molecular motion decreases and temperatures drop, less energy is present to break the hydrogen bonds between water molecules. These bonds remain intact and begin to form a rigid, lattice-like structure (e.g., ice) (Figure \(4\)a). When frozen, ice is less dense than liquid water (the molecules are farther apart). This means that ice floats on the surface of a body of water (Figure \(4\)b). In lakes, ponds, and oceans, ice will form on the surface of the water, creating an insulating barrier to protect the animal and plant life beneath from freezing in the water. If this did not happen, plants and animals living in water would freeze in a block of ice and could not move freely, making life in cold temperatures difficult or impossible.
Water Is an Excellent Solvent
Because water is polar, with slight positive and negative charges, ionic compounds and polar molecules can readily dissolve in it. Water is, therefore, what is referred to as a solvent—a substance capable of dissolving another substance. The charged particles will form hydrogen bonds with a surrounding layer of water molecules. This is referred to as a sphere of hydration and serves to keep the particles separated or dispersed in the water. In the case of table salt (NaCl) mixed in water (Figure \(5\)), the sodium and chloride ions separate, or dissociate, in the water, and spheres of hydration are formed around the ions. A positively charged sodium ion is surrounded by the partially negative charges of oxygen atoms in water molecules. A negatively charged chloride ion is surrounded by the partially positive charges of hydrogen atoms in water molecules. These spheres of hydration are also referred to as hydration shells. The polarity of the water molecule makes it an effective solvent and is important in its many roles in living systems.
Water Is Cohesive
Have you ever filled up a glass of water to the very top and then slowly added a few more drops? Before it overflows, the water actually forms a dome-like shape above the rim of the glass. This water can stay above the glass because of the property of cohesion. In cohesion, water molecules are attracted to each other (because of hydrogen bonding), keeping the molecules together at the liquid-air (gas) interface, although there is no more room in the glass. Cohesion gives rise to surface tension, the capacity of a substance to withstand rupture when placed under tension or stress. When you drop a small scrap of paper onto a droplet of water, the paper floats on top of the water droplet, although the object is denser (heavier) than the water. This occurs because of the surface tension that is created by the water molecules. Cohesion and surface tension keep the water molecules intact and the item floating on the top. It is even possible to “float” a steel needle on top of a glass of water if you place it gently, without breaking the surface tension (Figure \(6\)).
These cohesive forces are also related to the water’s property of adhesion, or the attraction between water molecules and other molecules. This is observed when water “climbs” up a straw placed in a glass of water. You will notice that the water appears to be higher on the sides of the straw than in the middle. This is because the water molecules are attracted to the straw and therefore adhere to it.
Cohesive and adhesive forces are important for sustaining life. For example, because of these forces, water can flow up from the roots to the tops of plants to feed the plant.
Query \(1\) | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/02%3A_Chemistry_for_Biology/2.03%3A_Water.txt |
The pH of a solution is a measure of its acidity or alkalinity. You have probably used litmus paper, paper that has been treated with a natural water-soluble dye so it can be used as a pH indicator, to test how much acid or base (alkalinity) exists in a solution. You might have even used some to make sure the water in an outdoor swimming pool is properly treated. In both cases, this pH test measures the amount of hydrogen ions that exists in a given solution. High concentrations of hydrogen ions yield a low pH, whereas low levels of hydrogen ions result in a high pH. The overall concentration of hydrogen ions is inversely related to its pH and can be measured on the pH scale (Figure \(1\)). Therefore, the more hydrogen ions present, the lower the pH; conversely, the fewer hydrogen ions, the higher the pH.
The pH scale ranges from 0 to 14. A change of one unit on the pH scale represents a change in the concentration of hydrogen ions by a factor of 10, a change in two units represents a change in the concentration of hydrogen ions by a factor of 100. Thus, small changes in pH represent large changes in the concentrations of hydrogen ions. Pure water is neutral. It is neither acidic nor basic, and has a pH of 7.0. Anything below 7.0 (ranging from 0.0 to 6.9) is acidic, and anything above 7.0 (from 7.1 to 14.0) is alkaline (basic). The blood in your veins is slightly alkaline (pH = 7.4). The environment in your stomach is highly acidic (pH = 1 to 2). Orange juice is mildly acidic (pH = approximately 3.5), whereas baking soda is basic (pH = 9.0).
Acids are substances that provide hydrogen ions (H+) and lower pH, whereas bases provide hydroxide ions (OH) and raise pH. The stronger the acid, the more readily it donates H+. For example, hydrochloric acid and lemon juice are very acidic and readily give up H+ when added to water. Conversely, bases are those substances that readily donate OH. The OH ions combine with H+ to produce water, which raises a substance’s pH. Sodium hydroxide and many household cleaners are very alkaline and give up OH rapidly when placed in water, thereby raising the pH.
Most cells in our bodies operate within a very narrow window of the pH scale, typically ranging only from 7.2 to 7.6. If the pH of the body is outside of this range, the respiratory system malfunctions, as do other organs in the body. Cells no longer function properly, and proteins will break down. Deviation outside of the pH range can induce coma or even cause death.
So how is it that we can ingest or inhale acidic or basic substances and not die? Buffers are the key. Buffers readily absorb excess H+ or OH, keeping the pH of the body carefully maintained in the aforementioned narrow range. Carbon dioxide is part of a prominent buffer system in the human body; it keeps the pH within the proper range. This buffer system involves carbonic acid (H2CO3) and bicarbonate (HCO3) anion. If too much H+ enters the body, bicarbonate will combine with the H+ to create carbonic acid and limit the decrease in pH. Likewise, if too much OH is introduced into the system, carbonic acid will rapidly dissociate into bicarbonate and H+ ions. The H+ ions can combine with the OH ions, limiting the increase in pH. While carbonic acid is an important product in this reaction, its presence is fleeting because the carbonic acid is released from the body as carbon dioxide gas each time we breathe. Without this buffer system, the pH in our bodies would fluctuate too much and we would fail to survive.
Query \(1\)
Query \(2\)
2.05: Absolutely Necessary Chemistry Summary
Matter
• Matter is anything that occupies space and has mass.
• Matter is made up of atoms of different elements.
• All of the 92 elements that occur naturally have unique qualities that allow them to combine in various ways to create compounds or molecules.
• Atoms consist of protons, neutrons, and electrons.
• Atoms are the smallest units of an element that retain all of the properties of that element.
Chemical Bonds
• Electrons can be donated or shared between atoms to create bonds.
• Ionic bonds form between a positively and a negatively charged atom. They are fairly strong bonds.
• Covalent bonds form when atoms share one or more electrons. They are very strong bonds.
• Hydrogen bonds form between partially charged atoms. They are weak bonds.
• van der Waals interactions form between polar, covalently bound atoms. They are weak attractions that are often temporary.
Water
• is POLAR, allowing for the formation of hydrogen bonds,
• is an excellent SOLVENT: because water is polar, it allows ions and other polar molecules to dissolve.
• STABILIZES TEMPERATURE: the hydrogen bonds between water molecules give water the ability to hold heat better than many other substances. As the temperature rises, the hydrogen bonds between water continually break and reform, allowing for the overall temperature to remain stable, although increased energy is added to the system.
• is COHESIVE: hydrogen bonds allow for the property of surface tension.
pH, Acids, Bases, and Buffers
• The pH of a solution is a measure of the concentration of hydrogen ions in the solution. The pH scale ranges from 0 to 14.
• A solution with an equal number of hydrogen ions and hydroxide ions is neutral and has a pH of 7.
• A solution with a high number of hydrogen ions is acidic and has a low pH value (below 7).
• A solution with a high number of hydroxide ions is basic and has a high pH value (above 7).
• Buffers are solutions that moderate pH changes when an acid or base is added to the buffer system. Buffers are important in biological systems because of their ability to maintain constant pH conditions. | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/02%3A_Chemistry_for_Biology/2.04%3A_Buffers_pH_Acids_and_Bases.txt |
Learning Outcomes
• Describe the structure of biologically-important molecules (carbohydrates, lipids, proteins, nucleic acids, water) and how their structure leads to their function.
Food provides an organism with nutrients—the matter it needs to survive. Many of these critical nutrients come in the form of biological macromolecules, or large molecules necessary for life. These macromolecules are built from different combinations of smaller organic molecules. What specific types of biological macromolecules do living things require? How are these molecules formed? What functions do they serve? In this chapter, we will explore these questions.
There are four major classes of biological macromolecules (carbohydrates, lipids, proteins, and nucleic acids), and each is an important component of the cell and performs a wide array of functions. Combined, these molecules make up the majority of a cell’s mass. Biological macromolecules are organic, meaning that they contain carbon atoms. In addition, they may contain atoms of hydrogen, oxygen, nitrogen, phosphorus, sulfur, and additional minor elements.
These molecules are made up of subunits called monomers. Each type of biological molecule is made up of different monomers. The monomers are connected together into a chain by strong covalent bonds. It is important that covalent bonds connect the monomers. If they were connected by hydrogen bonds the monomers would easily separate from each other and the biological molecule would come apart. If ionic bonds connected the monomers, the biological molecule would be likely to fall apart if it came into contact with water.
Query \(1\)
03: Biological Molecules
Carbohydrates are macromolecules with which most consumers are somewhat familiar. To lose weight, some individuals adhere to “low-carb” diets. Athletes, in contrast, often “carb-load” before important competitions to ensure that they have sufficient energy to compete at a high level. Carbohydrates are, in fact, an essential part of our diet; grains, fruits, and vegetables are all natural sources of carbohydrates. Carbohydrates provide energy to the body, particularly through glucose, a simple sugar. Carbohydrates also have other important functions in humans, animals, and plants.
Carbohydrates can be represented by the stoichiometric formula (CH2O)n, where n is the number of carbons in the molecule. In other words, the ratio of carbon to hydrogen to oxygen is 1:2:1 in carbohydrate molecules. This formula also explains the origin of the term “carbohydrate”: the components are carbon (“carbo”) and the components of water (hence, “hydrate”). Carbohydrates are classified into three subtypes: monosaccharides, disaccharides, and polysaccharides.
Monosaccharides
Monosaccharides (mono- = “one”; sacchar- = “sweet”) are simple sugars, the most common of which is glucose. In monosaccharides, the number of carbons usually ranges from three to seven. Most monosaccharide names end with the suffix -ose.
The chemical formula for glucose is C6H12O6. In humans, glucose is an important source of energy. During cellular respiration, energy is released from glucose, and that energy is used to help make adenosine triphosphate (ATP). Plants synthesize glucose using carbon dioxide and water, and glucose in turn is used for energy requirements for the plant. Excess glucose is often stored as starch that is catabolized (the breakdown of larger molecules by cells) by humans and other animals that feed on plants.
Galactose (part of lactose, or milk sugar) and fructose (found in sucrose, in fruit) are other common monosaccharides. Although glucose, galactose, and fructose all have the same chemical formula (C6H12O6), they differ structurally and chemically (and are known as isomers) because of the different arrangement of functional groups around the asymmetric carbon; all of these monosaccharides have more than one asymmetric carbon. Within one monosaccharide, all of the atoms are connected to each other with strong covalent bonds.
Disaccharides
Disaccharides (di- = “two”) form when two monosaccharides undergo a dehydration reaction (also known as a condensation reaction or dehydration synthesis). During this process, the hydroxyl (OH) group of one monosaccharide combines with the hydrogen of another monosaccharide, releasing a molecule of water and forming a covalent bond which joins the two monosaccharides together.
Common disaccharides include lactose, maltose, and sucrose (Figure \(3\)). Lactose is a disaccharide consisting of the monomers glucose and galactose. It is formed by a dehydration reaction between the glucose and the galactose molecules, which removes a water molecule and forms a covalent bond. It is found naturally in milk. Maltose, or malt sugar, is a disaccharide composed of two glucose molecules connected by a covalent bond. The most common disaccharide is sucrose, or table sugar, which is composed of the monomers glucose and fructose, also connected by a covalent bond.
Polysaccharides
A long chain of monosaccharides linked by glycosidic bonds is known as a polysaccharide (poly- = “many”). The chain may be branched or unbranched, and it may contain different types of monosaccharides. All of the monosaccharides are connected together by covalent bonds. The molecular weight may be 100,000 daltons or more depending on the number of monomers joined. Starch, glycogen, cellulose, and chitin are primary examples of polysaccharides.
Starch is the stored form of sugars in plants and is made up of a mixture of amylose and amylopectin (both polymers of glucose). Basically, starch is a long chain of glucose monomers. Plants are able to synthesize glucose, and the excess glucose, beyond the plant’s immediate energy needs, is stored as starch in different plant parts, including roots and seeds. The starch in the seeds provides food for the embryo as it germinates and can also act as a source of food for humans and animals. The starch that is consumed by humans is broken down by enzymes, such as salivary amylases, into smaller molecules, such as maltose and glucose. The cells can then absorb the glucose.
Glycogen is the storage form of glucose in humans and other vertebrates and is made up of monomers of glucose. Glycogen is the animal equivalent of starch and is a highly branched molecule usually stored in liver and muscle cells. Whenever blood glucose levels decrease, glycogen is broken down to release glucose in a process known as glycogenolysis.
Cellulose is the most abundant natural biopolymer. The cell wall of plants is mostly made of cellulose; this provides structural support to the cell. Wood and paper are mostly cellulosic in nature. Cellulose is made up of glucose monomers (Figure \(5\)).
Carbohydrates serve various functions in different animals. Arthropods (insects, crustaceans, and others) have an outer skeleton, called the exoskeleton, which protects their internal body parts (as seen in the bee in Figure \(6\)). This exoskeleton is made of the biological macromolecule chitin, which is a polysaccharide-containing nitrogen. It is made of repeating units of N-acetyl-β-d-glucosamine, a modified sugar. Chitin is also a major component of fungal cell walls; fungi are neither animals nor plants and form a kingdom of their own in the domain Eukarya.
How does carbohydrate structure relate to function?
Energy can be stored within the bonds of a molecule. Bonds connecting two carbon atoms or connecting a carbon atom to a hydrogen atom are high energy bonds. Breaking these bonds releases energy. This is why our cells can get energy from a molecule of glucose (C6H12O6).
Polysaccharides form long, fibrous chains which are able to build strong structures such as cell walls. | textbooks/bio/Introductory_and_General_Biology/Principles_of_Biology/01%3A_Chapter_1/03%3A_Biological_Molecules/3.01%3A_Carbohydrates.txt |
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