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Skills to Develop
• Name and describe the distinguishing features of the three main groups of mammals
• Describe the proposed line of descent that produced mammals
• List some derived features that may have arisen in response to mammals’ need for constant, high-level metabolism
Mammals are vertebrates that possess hair and mammary glands. Several other characteristics are distinctive to mammals, including certain features of the jaw, skeleton, integument, and internal anatomy. Modern mammals belong to three clades: monotremes, marsupials, and eutherians (or placental mammals).
Characteristics of Mammals
The presence of hair is one of the most obvious signs of a mammal. Although it is not very extensive on certain species, such as whales, hair has many important functions for mammals. Mammals are endothermic, and hair provides insulation to retain heat generated by metabolic work. Hair traps a layer of air close to the body, retaining heat. Along with insulation, hair can serve as a sensory mechanism via specialized hairs called vibrissae, better known as whiskers. These attach to nerves that transmit information about sensation, which is particularly useful to nocturnal or burrowing mammals. Hair can also provide protective coloration or be part of social signaling, such as when an animal’s hair stands “on end.”
Mammalian integument, or skin, includes secretory glands with various functions. Sebaceous glands produce a lipid mixture called sebum that is secreted onto the hair and skin for water resistance and lubrication. Sebaceous glands are located over most of the body. Eccrine glands produce sweat, or perspiration, which is mainly composed of water. In most mammals, eccrine glands are limited to certain areas of the body, and some mammals do not possess them at all. However, in primates, especially humans, sweat figures prominently in thermoregulation, regulating the body through evaporative cooling. Sweat glands are located over most of the body surface in primates. Apocrine glands, or scent glands, secrete substances that are used for chemical communication, such as in skunks. Mammary glands produce milk that is used to feed newborns. While male monotremes and eutherians possess mammary glands, male marsupials do not. Mammary glands likely are modified sebaceous or eccrine glands, but their evolutionary origin is not entirely clear.
The skeletal system of mammals possesses many unique features. The lower jaw of mammals consists of only one bone, the dentary. The jaws of other vertebrates are composed of more than one bone. In mammals, the dentary bone joins the skull at the squamosal bone, while in other vertebrates, the quadrate bone of the jaw joins with the articular bone of the skull. These bones are present in mammals, but they have been modified to function in hearing and form bones in the middle ear (Figure \(1\)). Other vertebrates possess only one middle ear bone, the stapes. Mammals have three: the malleus, incus, and stapes. The malleus originated from the articular bone, whereas the incus originated from the quadrate bone. This arrangement of jaw and ear bones aids in distinguishing fossil mammals from fossils of other synapsids.
The adductor muscle that closes the jaw is composed of two muscles in mammals: the temporalis and the masseter. These allow side-to-side movement of the jaw, making chewing possible, which is unique to mammals. Most mammals have heterodont teeth, meaning that they have different types and shapes of teeth rather than just one type and shape of tooth. Most mammals are diphyodonts, meaning that they have two sets of teeth in their lifetime: deciduous or “baby” teeth, and permanent teeth. Other vertebrates are polyphyodonts, that is, their teeth are replaced throughout their entire life.
Mammals, like birds, possess a four-chambered heart. Mammals also have a specialized group of cardiac fibers located in the walls of their right atrium called the sinoatrial node, or pacemaker, which determines the rate at which the heart beats. Mammalian erythrocytes (red blood cells) do not have nuclei, whereas the erythrocytes of other vertebrates are nucleated.
The kidneys of mammals have a portion of the nephron called the loop of Henle or nephritic loop, which allows mammals to produce urine with a high concentration of solutes, higher than that of the blood. Mammals lack a renal portal system, which is a system of veins that moves blood from the hind or lower limbs and region of the tail to the kidneys. Renal portal systems are present in all other vertebrates except jawless fishes. A urinary bladder is present in all mammals.
Mammalian brains have certain characteristics that differ from other vertebrates. In some, but not all mammals, the cerebral cortex, the outermost part of the cerebrum, is highly folded, allowing for a greater surface area than is possible with a smooth cortex. The optic lobes, located in the midbrain, are divided into two parts in mammals, whereas other vertebrates possess a single, undivided lobe. Eutherian mammals also possess a specialized structure that links the two cerebral hemispheres, called the corpus callosum.
Evolution of Mammals
Mammals are synapsids, meaning they have a single opening in the skull. They are the only living synapsids, as earlier forms became extinct by the Jurassic period. The early non-mammalian synapsids can be divided into two groups, the pelycosaurs and the therapsids. Within the therapsids, a group called the cynodonts are thought to be the ancestors of mammals (Figure \(2\)).
A key characteristic of synapsids is endothermy, rather than the ectothermy seen in most other vertebrates. The increased metabolic rate required to internally modify body temperature went hand in hand with changes to certain skeletal structures. The later synapsids, which had more evolved characteristics unique to mammals, possess cheeks for holding food and heterodont teeth, which are specialized for chewing, mechanically breaking down food to speed digestion and releasing the energy needed to produce heat. Chewing also requires the ability to chew and breathe at the same time, which is facilitated by the presence of a secondary palate. A secondary palate separates the area of the mouth where chewing occurs from the area above where respiration occurs, allowing breathing to proceed uninterrupted during chewing. A secondary palate is not found in pelycosaurs but is present in cynodonts and mammals. The jawbone also shows changes from early synapsids to later ones. The zygomatic arch, or cheekbone, is present in mammals and advanced therapsids such as cynodonts, but is not present in pelycosaurs. The presence of the zygomatic arch suggests the presence of the masseter muscle, which closes the jaw and functions in chewing.
In the appendicular skeleton, the shoulder girdle of therian mammals is modified from that of other vertebrates in that it does not possess a procoracoid bone or an interclavicle, and the scapula is the dominant bone.
Mammals evolved from therapsids in the late Triassic period, as the earliest known mammal fossils are from the early Jurassic period, some 205 million years ago. Early mammals were small, about the size of a small rodent. Mammals first began to diversify in the Mesozoic Era, from the Jurassic to the Cretaceous periods, although most of these mammals were extinct by the end of the Mesozoic. During the Cretaceous period, another radiation of mammals began and continued through the Cenozoic Era, about 65 million years ago.
Living Mammals
The eutherians, or placental mammals, and the marsupials together comprise the clade of therian mammals. Monotremes, or metatherians, form their sister clade.
There are three living species of monotremes: the platypus and two species of echidnas, or spiny anteaters. The leathery-beaked platypus belongs to the family Ornithorhynchidae (“bird beak”), whereas echidnas belong to the family Tachyglossidae (“sticky tongue”) (Figure \(3\)). The platypus and one species of echidna are found in Australia, and the other species of echidna is found in New Guinea. Monotremes are unique among mammals as they lay eggs, rather than giving birth to live young. The shells of their eggs are not like the hard shells of birds, but are a leathery shell, similar to the shells of reptile eggs. Monotremes have no teeth.
Marsupials are found primarily in Australia, though the opossum is found in North America. Australian marsupials include the kangaroo, koala, bandicoot, Tasmanian devil (Figure \(4\)), and several other species. Most species of marsupials possess a pouch in which the very premature young reside after birth, receiving milk and continuing to develop. Marsupials differ from eutherians in that there is a less complex placental connection: The young are born at an extremely early age and latch onto the nipple within the pouch.
Eutherians are the most widespread of the mammals, occurring throughout the world. There are 18 to 20 orders of placental mammals. Some examples are Insectivora, the insect eaters; Edentata, the toothless anteaters; Rodentia, the rodents; Cetacea, the aquatic mammals including whales; Carnivora, carnivorous mammals including dogs, cats, and bears; and Primates, which includes humans. Eutherian mammals are sometimes called placental mammals because all species possess a complex placenta that connects a fetus to the mother, allowing for gas, fluid, and nutrient exchange. While other mammals possess a less complex placenta or briefly have a placenta, all eutherians possess a complex placenta during gestation.
Summary
Mammals in general are vertebrates that possess hair and mammary glands. The mammalian integument includes various secretory glands, including sebaceous glands, eccrine glands, apocrine glands, and mammary glands. Mammals are synapsids, meaning that they have a single opening in the skull. A key characteristic of synapsids is endothermy rather than the ectothermy seen in other vertebrates. Mammals probably evolved from therapsids in the late Triassic period, as the earliest known mammal fossils are from the early Jurassic period. There are three groups of mammals living today: monotremes, marsupials, and eutherians. Monotremes are unique among mammals as they lay eggs, rather than giving birth to young. Eutherian mammals are sometimes called placental mammals, because all species possess a complex placenta that connects a fetus to the mother, allowing for gas, fluid, and nutrient exchange.
Glossary
apocrine gland
scent gland that secretes substances that are used for chemical communication
dentary
single bone that comprises the lower jaw of mammals
diphyodont
refers to the possession of two sets of teeth in a lifetime
eccrine gland
sweat gland
eutherian mammal
mammal that possesses a complex placenta, which connects a fetus to the mother; sometimes called placental mammals
heterodont tooth
different types of teeth that are modified for different purposes
mammal
one of the groups of endothermic vertebrates that possesses hair and mammary glands
mammary gland
in female mammals, a gland that produces milk for newborns
marsupial
one of the groups of mammals that includes the kangaroo, koala, bandicoot, Tasmanian devil, and several other species; young develop within a pouch
monotreme
egg-laying mammal
Ornithorhynchidae
clade that includes the duck-billed platypus
sebaceous gland
in mammals, a skin gland that produce a lipid mixture called sebum
Tachyglossidae
clade that includes the echidna or spiny anteater | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/5%3A_Biological_Diversity/29%3A_Vertebrates/29.6%3A_Mammals.txt |
Skills to Develop
• Describe the derived features that distinguish primates from other animals
• Explain why scientists are having difficulty determining the true lines of descent in hominids
Order Primates of class Mammalia includes lemurs, tarsiers, monkeys, apes, and humans. Non-human primates live primarily in the tropical or subtropical regions of South America, Africa, and Asia. They range in size from the mouse lemur at 30 grams (1 ounce) to the mountain gorilla at 200 kilograms (441 pounds). The characteristics and evolution of primates is of particular interest to us as it allows us to understand the evolution of our own species.
Characteristics of Primates
All primate species possess adaptations for climbing trees, as they all descended from tree-dwellers. This arboreal heritage of primates has resulted in hands and feet that are adapted for brachiation, or climbing and swinging through trees. These adaptations include, but are not limited to: 1) a rotating shoulder joint, 2) a big toe that is widely separated from the other toes and thumbs, which are widely separated from fingers (except humans), which allow for gripping branches, 3) stereoscopic vision, two overlapping fields of vision from the eyes, which allows for the perception of depth and gauging distance. Other characteristics of primates are brains that are larger than those of most other mammals, claws that have been modified into flattened nails, typically only one offspring per pregnancy, and a trend toward holding the body upright.
Order Primates is divided into two groups: prosimians and anthropoids. Prosimians include the bush babies of Africa, the lemurs of Madagascar, and the lorises, pottos, and tarsiers of Southeast Asia. Anthropoids include monkeys, apes, and humans. In general, prosimians tend to be nocturnal (in contrast to diurnal anthropoids) and exhibit a smaller size and smaller brain than anthropoids.
Evolution of Primates
The first primate-like mammals are referred to as proto-primates. They were roughly similar to squirrels and tree shrews in size and appearance. The existing fossil evidence (mostly from North Africa) is very fragmented. These proto-primates remain largely mysterious creatures until more fossil evidence becomes available. The oldest known primate-like mammals with a relatively robust fossil record is Plesiadapis (although some researchers do not agree that Plesiadapis was a proto-primate). Fossils of this primate have been dated to approximately 55 million years ago. Plesiadapiforms were proto-primates that had some features of the teeth and skeleton in common with true primates. They were found in North America and Europe in the Cenozoic and went extinct by the end of the Eocene.
The first true primates were found in North America, Europe, Asia, and Africa in the Eocene Epoch. These early primates resembled present-day prosimians such as lemurs. Evolutionary changes continued in these early primates, with larger brains and eyes, and smaller muzzles being the trend. By the end of the Eocene Epoch, many of the early prosimian species went extinct due either to cooler temperatures or competition from the first monkeys.
Anthropoid monkeys evolved from prosimians during the Oligocene Epoch. By 40 million years ago, evidence indicates that monkeys were present in the New World (South America) and the Old World (Africa and Asia). New World monkeys are also called Platyrrhini—a reference to their broad noses (Figure \(1\)). Old World monkeys are called Catarrhini—a reference to their narrow noses. There is still quite a bit of uncertainty about the origins of the New World monkeys. At the time the platyrrhines arose, the continents of South American and Africa had drifted apart. Therefore, it is thought that monkeys arose in the Old World and reached the New World either by drifting on log rafts or by crossing land bridges. Due to this reproductive isolation, New World monkeys and Old World monkeys underwent separate adaptive radiations over millions of years. The New World monkeys are all arboreal, whereas Old World monkeys include arboreal and ground-dwelling species.
Apes evolved from the catarrhines in Africa midway through the Cenozoic, approximately 25 million years ago. Apes are generally larger than monkeys and they do not possess a tail. All apes are capable of moving through trees, although many species spend most their time on the ground. Apes are more intelligent than monkeys, and they have relatively larger brains proportionate to body size. The apes are divided into two groups. The lesser apes comprise the family Hylobatidae, including gibbons and siamangs. The great apes include the genera Pan (chimpanzees and bonobos) (Figure \(2\)a), Gorilla (gorillas), Pongo (orangutans), and Homo (humans) (Figure \(2\)b). The very arboreal gibbons are smaller than the great apes; they have low sexual dimorphism (that is, the genders are not markedly different in size); and they have relatively longer arms used for swinging through trees.
Human Evolution
The family Hominidae of order Primates includes the hominoids: the great apes (Figure \(3\)). Evidence from the fossil record and from a comparison of human and chimpanzee DNA suggests that humans and chimpanzees diverged from a common hominoid ancestor approximately 6 million years ago. Several species evolved from the evolutionary branch that includes humans, although our species is the only surviving member. The term hominin is used to refer to those species that evolved after this split of the primate line, thereby designating species that are more closely related to humans than to chimpanzees. Hominins were predominantly bipedal and include those groups that likely gave rise to our species—including Australopithecus, Homo habilis, and Homo erectus—and those non-ancestral groups that can be considered “cousins” of modern humans, such as Neanderthals. Determining the true lines of descent in hominins is difficult. In years past, when relatively few hominin fossils had been recovered, some scientists believed that considering them in order, from oldest to youngest, would demonstrate the course of evolution from early hominins to modern humans. In the past several years, however, many new fossils have been found, and it is clear that there was often more than one species alive at any one time and that many of the fossils found (and species named) represent hominin species that died out and are not ancestral to modern humans.
Very Early Hominins
Three species of very early hominids have made news in the past few years. The oldest of these, Sahelanthropus tchadensis, has been dated to nearly 7 million years ago. There is a single specimen of this genus, a skull that was a surface find in Chad. The fossil, informally called “Toumai,” is a mosaic of primitive and evolved characteristics, and it is unclear how this fossil fits with the picture given by molecular data, namely that the line leading to modern humans and modern chimpanzees apparently bifurcated about 6 million years ago. It is not thought at this time that this species was an ancestor of modern humans.
A second, younger species, Orrorin tugenensis, is also a relatively recent discovery, found in 2000. There are several specimens of Orrorin. It is not known whether Orrorin was a human ancestor, but this possibility has not been ruled out. Some features of Orrorin are more similar to those of modern humans than are the australopiths, although Orrorin is much older.
A third genus, Ardipithecus, was discovered in the 1990s, and the scientists who discovered the first fossil found that some other scientists did not believe the organism to be a biped (thus, it would not be considered a hominid). In the intervening years, several more specimens of Ardipithecus, classified as two different species, demonstrated that the organism was bipedal. Again, the status of this genus as a human ancestor is uncertain.
Early Hominins: Genus Australopithecus
Australopithecus (“southern ape”) is a genus of hominin that evolved in eastern Africa approximately 4 million years ago and went extinct about 2 million years ago. This genus is of particular interest to us as it is thought that our genus, genus Homo, evolved from Australopithecus about 2 million years ago (after likely passing through some transitional states). Australopithecus had a number of characteristics that were more similar to the great apes than to modern humans. For example, sexual dimorphism was more exaggerated than in modern humans. Males were up to 50 percent larger than females, a ratio that is similar to that seen in modern gorillas and orangutans. In contrast, modern human males are approximately 15 to 20 percent larger than females. The brain size of Australopithecus relative to its body mass was also smaller than modern humans and more similar to that seen in the great apes. A key feature that Australopithecus had in common with modern humans was bipedalism, although it is likely that Australopithecus also spent time in trees. Hominin footprints, similar to those of modern humans, were found in Laetoli, Tanzania and dated to 3.6 million years ago. They showed that hominins at the time of Australopithecus were walking upright.
There were a number of Australopithecus species, which are often referred to as australopiths. Australopithecus anamensis lived about 4.2 million years ago. More is known about another early species, Australopithecus afarensis, which lived between 3.9 and 2.9 million years ago. This species demonstrates a trend in human evolution: the reduction of the dentition and jaw in size. A. afarensis (Figure \(4\)) had smaller canines and molars compared to apes, but these were larger than those of modern humans. Its brain size was 380–450 cubic centimeters, approximately the size of a modern chimpanzee brain. It also had prognathic jaws, which is a relatively longer jaw than that of modern humans. In the mid-1970s, the fossil of an adult female A. afarensis was found in the Afar region of Ethiopia and dated to 3.24 million years ago (Figure \(5\)). The fossil, which is informally called “Lucy,” is significant because it was the most complete australopith fossil found, with 40 percent of the skeleton recovered.
Australopithecus africanus lived between 2 and 3 million years ago. It had a slender build and was bipedal, but had robust arm bones and, like other early hominids, may have spent significant time in trees. Its brain was larger than that of A. afarensis at 500 cubic centimeters, which is slightly less than one-third the size of modern human brains. Two other species, Australopithecus bahrelghazali and Australopithecus garhi, have been added to the roster of australopiths in recent years.
A Dead End: Genus Paranthropus
The australopiths had a relatively slender build and teeth that were suited for soft food. In the past several years, fossils of hominids of a different body type have been found and dated to approximately 2.5 million years ago. These hominids, of the genus Paranthropus, were relatively large and had large grinding teeth. Their molars showed heavy wear, suggesting that they had a coarse and fibrous vegetarian diet as opposed to the partially carnivorous diet of the australopiths. Paranthropus includes Paranthropus robustus of South Africa, and Paranthropus aethiopicus and Paranthropus boisei of East Africa. The hominids in this genus went extinct more than 1 million years ago and are not thought to be ancestral to modern humans, but rather members of an evolutionary branch on the hominin tree that left no descendants.
Early Hominins: Genus Homo
The human genus, Homo, first appeared between 2.5 and 3 million years ago. For many years, fossils of a species called H. habilis were the oldest examples in the genus Homo, but in 2010, a new species called Homo gautengensis was discovered and may be older. Compared to A. africanus, H. habilis had a number of features more similar to modern humans. H. habilis had a jaw that was less prognathic than the australopiths and a larger brain, at 600–750 cubic centimeters. However, H. habilis retained some features of older hominin species, such as long arms. The name H. habilis means “handy man,” which is a reference to the stone tools that have been found with its remains.
Link to Learning
Visit this site for a video about Smithsonian paleontologist Briana Pobiner explaining the link between hominin eating of meat and evolutionary trends.
H. erectus appeared approximately 1.8 million years ago (Figure \(6\)). It is believed to have originated in East Africa and was the first hominin species to migrate out of Africa. Fossils of H. erectus have been found in India, China, Java, and Europe, and were known in the past as “Java Man” or “Peking Man.” H. erectus had a number of features that were more similar to modern humans than those of H. habilis. H. erectus was larger in size than earlier hominins, reaching heights up to 1.85 meters and weighing up to 65 kilograms, which are sizes similar to those of modern humans. Its degree of sexual dimorphism was less than earlier species, with males being 20 to 30 percent larger than females, which is close to the size difference seen in our species. H. erectus had a larger brain than earlier species at 775–1,100 cubic centimeters, which compares to the 1,130–1,260 cubic centimeters seen in modern human brains. H. erectus also had a nose with downward-facing nostrils similar to modern humans, rather than the forward facing nostrils found in other primates. Longer, downward-facing nostrils allow for the warming of cold air before it enters the lungs and may have been an adaptation to colder climates. Artifacts found with fossils of H. erectus suggest that it was the first hominin to use fire, hunt, and have a home base. H. erectus is generally thought to have lived until about 50,000 years ago.
Humans: Homo sapiens
A number of species, sometimes called archaic Homo sapiens, apparently evolved from H. erectus starting about 500,000 years ago. These species include Homo heidelbergensis, Homo rhodesiensis, and Homo neanderthalensis. These archaic H. sapiens had a brain size similar to that of modern humans, averaging 1,200–1,400 cubic centimeters. They differed from modern humans by having a thick skull, a prominent brow ridge, and a receding chin. Some of these species survived until 30,000–10,000 years ago, overlapping with modern humans (Figure \(7\)).
There is considerable debate about the origins of anatomically modern humans or Homo sapiens sapiens. As discussed earlier, H. erectus migrated out of Africa and into Asia and Europe in the first major wave of migration about 1.5 million years ago. It is thought that modern humans arose in Africa from H. erectus and migrated out of Africa about 100,000 years ago in a second major migration wave. Then, modern humans replaced H. erectus species that had migrated into Asia and Europe in the first wave.
This evolutionary timeline is supported by molecular evidence. One approach to studying the origins of modern humans is to examine mitochondrial DNA (mtDNA) from populations around the world. Because a fetus develops from an egg containing its mother’s mitochondria (which have their own, non-nuclear DNA), mtDNA is passed entirely through the maternal line. Mutations in mtDNA can now be used to estimate the timeline of genetic divergence. The resulting evidence suggests that all modern humans have mtDNA inherited from a common ancestor that lived in Africa about 160,000 years ago. Another approach to the molecular understanding of human evolution is to examine the Y chromosome, which is passed from father to son. This evidence suggests that all men today inherited a Y chromosome from a male that lived in Africa about 140,000 years ago.
Summary
All primate species possess adaptations for climbing trees, as they all probably descended from tree-dwellers, although not all species are arboreal. Other characteristics of primates are brains that are larger than those of other mammals, claws that have been modified into flattened nails, typically only one young per pregnancy, stereoscopic vision, and a trend toward holding the body upright. Primates are divided into two groups: prosimians and anthropoids. Monkeys evolved from prosimians during the Oligocene Epoch. Apes evolved from catarrhines in Africa during the Miocene Epoch. Apes are divided into the lesser apes and the greater apes. Hominins include those groups that gave rise to our species, such as Australopithecus and H. erectus, and those groups that can be considered “cousins” of humans, such as Neanderthals. Fossil evidence shows that hominins at the time of Australopithecus were walking upright, the first evidence of bipedal hominins. A number of species, sometimes called archaic H. sapiens, evolved from H. erectus approximately 500,000 years ago. There is considerable debate about the origins of anatomically modern humans or H. sapiens sapiens.
Glossary
anthropoid
monkeys, apes, and humans
Australopithecus
genus of hominins that evolved in eastern Africa approximately 4 million years ago
brachiation
movement through trees branches via suspension from the arms
Catarrhini
clade of Old World monkeys
Gorilla
genus of gorillas
hominin
species that are more closely related to humans than chimpanzees
hominoid
pertaining to great apes and humans
Homo
genus of humans
Homo sapiens sapiens
anatomically modern humans
Hylobatidae
family of gibbons
Pan
genus of chimpanzees and bonobos
Platyrrhini
clade of New World monkeys
Plesiadapis
oldest known primate-like mammal
Pongo
genus of orangutans
Primates
order of lemurs, tarsiers, monkeys, apes, and humans
prognathic jaw
long jaw
prosimian
division of primates that includes bush babies of Africa, lemurs of Madagascar, and lorises, pottos, and tarsiers of Southeast Asia
stereoscopic vision
two overlapping fields of vision from the eyes that produces depth perception | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/5%3A_Biological_Diversity/29%3A_Vertebrates/29.7%3A_The_Evolution_of_Primates.txt |
29.1: Chordates
Review Questions
Which of the following is not contained in phylum Chordata?
1. Cephalochordata
2. Echinodermata
3. Urochordata
4. Vertebrata
Answer
B
Which group of invertebrates is most closely related to vertebrates?
1. cephalochordates
2. echinoderms
3. arthropods
4. urochordates
Answer
A
Free Response
What are the characteristic features of the chordates?
Answer
The characteristic features of the phylum Chordata are a notochord, a dorsal hollow nerve cord, pharyngeal slits, and a post-anal tail.
29.2: Fishes
Review Questions
Members of Chondrichthyes differ from members of Osteichthyes by having a ________.
1. jaw
2. bony skeleton
3. cartilaginous skeleton
4. two sets of paired fins
Answer
C
Members of Chondrichthyes are thought to be descended from fishes that had ________.
1. a cartilaginous skeleton
2. a bony skeleton
3. mucus glands
4. slime glands
Answer
B
Free Response
What can be inferred about the evolution of the cranium and vertebral column from examining hagfishes and lampreys?
Answer
Comparison of hagfishes with lampreys shows that the cranium evolved first in early vertebrates, as it is seen in hagfishes, which evolved earlier than lampreys. This was followed by evolution of the vertebral column, a primitive form of which is seen in lampreys and not in hagfishes.
Why did gnathostomes replace most agnathans?
Answer
Evolution of the jaw and paired fins permitted gnathostomes to diversify from the sedentary suspension feeding of agnathans to a mobile predatory lifestyle. The ability of gnathostomes to utilize new nutrient sources may be one reason why the gnathostomes replaced most agnathans.
29.3: Amphibians
Review Questions
Which of the following is not true of Acanthostega?
1. It was aquatic.
2. It had gills.
3. It had four limbs.
4. It laid shelled eggs.
Answer
D
Frogs belong to which order?
1. Anura
2. Urodela
3. Caudata
4. Apoda
Answer
A
Free Response
Explain why frogs are restricted to a moist environment.
Answer
A moist environment is required, as frog eggs lack a shell and dehydrate quickly in dry environments.
Describe the differences between the larval and adult stages of frogs.
Answer
The larval stage of frogs is the tadpole, which is usually a filter-feeding herbivore. Tadpoles usually have gills, a lateral line system, long-finned tails, and lack limbs. In the adult form, the gills and lateral line system disappear, and four limbs develop. The jaws grow larger, suitable for carnivorous feeding, and the digestive system transforms into the typical short gut of a predator. An eardrum and air-breathing lungs also develop.
29.4: Reptiles
Review Questions
During the Mesozoic period, diapsids diverged into_______.
1. pterosaurs and dinosaurs
2. mammals and reptiles
3. lepidosaurs and archosaurs
4. Testudines and Sphenodontia
Answer
C
Squamata includes_______.
1. crocodiles and alligators
2. turtles
3. tuataras
4. lizards and snakes
Answer
D
Free Response
Describe the functions of the three extra-embryonic membranes present in amniotic eggs.
Answer
The chorion facilitates the exchange of oxygen and carbon dioxide gases between the embryo and the surrounding air. The amnion protects the embryo from mechanical shock and prevents dehydration. The allantois stores nitrogenous wastes produced by the embryo and facilitates respiration.
What characteristics differentiate lizards and snakes?
Answer
Lizards differ from snakes by having eyelids, external ears, and less kinematic skulls.
29.5: Birds
Review Questions
A bird or feathered dinosaur is ________.
1. Neornithes
2. Archaeopteryx
3. Enantiornithes
4. Paleognathae
Answer
B
Which of the following feather types helps to reduce drag produced by wind resistance during flight?
1. flight feathers
2. primary feathers
3. secondary feathers
4. contour feathers
Answer
D
Free Response
Explain why birds are thought to have evolved from theropod dinosaurs.
Answer
This is suggested by similarities observed between theropod fossils and birds, specifically in the design of the hip and wrist bones, as well as the presence of a furcula, or wishbone, formed by the fusing of the clavicles.
Describe three skeletal adaptations that allow for flight in birds.
Answer
The sternum of birds is larger than that of other vertebrates, which accommodates the force required for flapping. Another skeletal modification is the fusion of the clavicles, forming the furcula or wishbone. The furcula is flexible enough to bend during flapping and provides support to the shoulder girdle during flapping. Birds also have pneumatic bones that are hollow rather than filled with tissue.
29.6: Mammals
Review Questions
Eccrine glands produce ________.
1. sweat
2. lipids
3. scents
4. milk
Answer
A
Monotremes include:
1. kangaroos
2. koalas
3. bandicoots
4. platypuses
Answer
D
Free Response
Describe three unique features of the mammalian skeletal system.
Answer
The lower jaw of mammals consists of only one bone, the dentary. The dentary bone joins the skull at the squamosal bone. Mammals have three bones of the middle ear. The adductor muscle that closes the jaw is composed of two muscles in mammals. Most mammals have heterodont teeth.
Describe three characteristics of the mammalian brain that differ from other vertebrates.
Answer
In some mammals, the cerebral cortex is highly folded, allowing for greater surface area than a smooth cortex. The optic lobes are divided into two parts in mammals. Eutherian mammals also possess a specialized structure that links the two cerebral hemispheres, called the corpus callosum.
29.7: The Evolution of Primates
Review Questions
Which of the following is not an anthropoid?
1. lemurs
2. monkeys
3. apes
4. humans
Answer
A
Which of the following is part of a clade believed to have died out, leaving no descendants?
1. Paranthropus robustus
2. Australopithecus africanus
3. Homo erectus
4. Homo sapiens sapiens
Answer
A
Free Response
How did archaic Homo sapiens differ from anatomically modern humans?
Answer
Archaic Homo sapiens differed from modern humans by having a thick skull and a prominent brow ridge, and lacking a prominent chin.
Why is it so difficult to determine the sequence of hominin ancestors that have led to modern Homo sapiens?
Answer
The immediate ancestors of humans were Australopithecus. All people past and present, along with the australopithecines, are hominins. We share the adaptation of being habitually bipedal. The earliest australopithecines very likely did not evolve until 5 million years ago. The primate fossil record for this crucial transitional period leading to australopithecines is still sketchy and somewhat confusing. By about 2.5 million years ago, there were at least two evolutionary lines of hominins descended from early australopithecines. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/5%3A_Biological_Diversity/29%3A_Vertebrates/29.E%3A_Vertebrates_%28Exercises%29.txt |
Like animals, plants contain cells with organelles in which specific metabolic activities take place. Unlike animals, however, plants use energy from sunlight to form sugars during photosynthesis. In addition, plant cells have cell walls, plastids, and a large central vacuole: structures that are not found in animal cells. Each of these cellular structures plays a specific role in plant structure and function.
• 30.0: Prelude to Plant Form and Physiology
While individual plant species are unique, all share a common structure: a plant body consisting of stems, roots, and leaves. They all transport water, minerals, and sugars produced through photosynthesis through the plant body in a similar manner. All plant species also respond to environmental factors, such as light, gravity, competition, temperature, and predation.
• 30.1: The Plant Body
Like animals, plants contain cells with organelles in which specific metabolic activities take place. Unlike animals, however, plants use energy from sunlight to form sugars during photosynthesis. In addition, plant cells have cell walls, plastids, and a large central vacuole: structures that are not found in animal cells. Each of these cellular structures plays a specific role in plant structure and function.
• 30.2: Stems
Plant stems, whether above or below ground, are characterized by the presence of nodes and internodes. Nodes are points of attachment for leaves, aerial roots, and flowers. The stem region between two nodes is called an internode. The stalk that extends from the stem to the base of the leaf is the petiole. An axillary bud is usually found in the axil—the area between the base of a leaf and the stem—where it can give rise to a branch or a flower.
• 30.3: Roots
The roots of seed plants have three major functions: anchoring the plant to the soil, absorbing water and minerals and transporting them upwards, and storing the products of photosynthesis. Some roots are modified to absorb moisture and exchange gases. Most roots are underground. Some plants, however, also have adventitious roots, which emerge above the ground from the shoot.
• 30.4: Leaves
Leaves are the main sites for photosynthesis: the process by which plants synthesize food. Most leaves are usually green, due to the presence of chlorophyll in the leaf cells. However, some leaves may have different colors, caused by other plant pigments that mask the green chlorophyll. The thickness, shape, and size of leaves are adapted to the environment. Each variation helps a plant species maximize its chances of survival in a particular habitat.
• 30.5: Transport of Water and Solutes in Plants
The structure of plant roots, stems, and leaves facilitates the transport of water, nutrients, and photosynthates throughout the plant. The phloem and xylem are the main tissues responsible for this movement. Water potential, evapotranspiration, and stomatal regulation influence how water and nutrients are transported in plants. To understand how these processes work, we must first understand the energetics of water potential.
• 30.6: Plant Sensory Systems and Responses
Animals can respond to environmental factors by moving to a new location. Plants, however, are rooted in place and must respond to the surrounding environmental factors. Plants have sophisticated systems to detect and respond to light, gravity, temperature, and physical touch. Receptors sense environmental factors and relay the information to effector systems—often through intermediate chemical messengers—to bring about plant responses.
• 30.E: Plant Form and Physiology (Exercises)
Thumbnail: Rose thorns. (CC BY 2.0; macrophile via Flickr).
30: Plant Form and Physiology
Plants are as essential to human existence as land, water, and air. Without plants, our day-to-day lives would be impossible because without oxygen from photosynthesis, aerobic life cannot be sustained. From providing food and shelter to serving as a source of medicines, oils, perfumes, and industrial products, plants provide humans with numerous valuable resources.
When you think of plants, most of the organisms that come to mind are vascular plants. These plants have tissues that conduct food and water, and they have seeds. Seed plants are divided into gymnosperms and angiosperms. Gymnosperms include the needle-leaved conifers—spruce, fir, and pine—as well as less familiar plants, such as ginkgos and cycads. Their seeds are not enclosed by a fleshy fruit. Angiosperms, also called flowering plants, constitute the majority of seed plants. They include broadleaved trees (such as maple, oak, and elm), vegetables (such as potatoes, lettuce, and carrots), grasses, and plants known for the beauty of their flowers (roses, irises, and daffodils, for example).
While individual plant species are unique, all share a common structure: a plant body consisting of stems, roots, and leaves. They all transport water, minerals, and sugars produced through photosynthesis through the plant body in a similar manner. All plant species also respond to environmental factors, such as light, gravity, competition, temperature, and predation. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.0%3A_Prelude_to_Plant_Form_and_Physiology.txt |
Skills to Develop
• Describe the shoot organ system and the root organ system
• Distinguish between meristematic tissue and permanent tissue
• Identify and describe the three regions where plant growth occurs
• Summarize the roles of dermal tissue, vascular tissue, and ground tissue
• Compare simple plant tissue with complex plant tissue
Like animals, plants contain cells with organelles in which specific metabolic activities take place. Unlike animals, however, plants use energy from sunlight to form sugars during photosynthesis. In addition, plant cells have cell walls, plastids, and a large central vacuole: structures that are not found in animal cells. Each of these cellular structures plays a specific role in plant structure and function.
Link to Learning
Watch Botany Without Borders, a video produced by the Botanical Society of America about the importance of plants.
Plant Organ Systems
In plants, just as in animals, similar cells working together form a tissue. When different types of tissues work together to perform a unique function, they form an organ; organs working together form organ systems. Vascular plants have two distinct organ systems: a shoot system, and a root system. The shoot system consists of two portions: the vegetative (non-reproductive) parts of the plant, such as the leaves and the stems, and the reproductive parts of the plant, which include flowers and fruits. The shoot system generally grows above ground, where it absorbs the light needed for photosynthesis. The root system, which supports the plants and absorbs water and minerals, is usually underground. Figure \(1\) shows the organ systems of a typical plant.
Plant Tissues
Plants are multicellular eukaryotes with tissue systems made of various cell types that carry out specific functions. Plant tissue systems fall into one of two general types: meristematic tissue, and permanent (or non-meristematic) tissue. Cells of the meristematic tissue are found in meristems, which are plant regions of continuous cell division and growth. Meristematic tissue cells are either undifferentiated or incompletely differentiated, and they continue to divide and contribute to the growth of the plant. In contrast, permanent tissue consists of plant cells that are no longer actively dividing.
Meristematic tissues consist of three types, based on their location in the plant. Apical meristems contain meristematic tissue located at the tips of stems and roots, which enable a plant to extend in length. Lateral meristems facilitate growth in thickness or girth in a maturing plant. Intercalary meristems occur only in monocots, at the bases of leaf blades and at nodes (the areas where leaves attach to a stem). This tissue enables the monocot leaf blade to increase in length from the leaf base; for example, it allows lawn grass leaves to elongate even after repeated mowing.
Meristems produce cells that quickly differentiate, or specialize, and become permanent tissue. Such cells take on specific roles and lose their ability to divide further. They differentiate into three main types: dermal, vascular, and ground tissue. Dermal tissue covers and protects the plant, and vascular tissue transports water, minerals, and sugars to different parts of the plant. Ground tissue serves as a site for photosynthesis, provides a supporting matrix for the vascular tissue, and helps to store water and sugars.
Secondary tissues are either simple (composed of similar cell types) or complex (composed of different cell types). Dermal tissue, for example, is a simple tissue that covers the outer surface of the plant and controls gas exchange. Vascular tissue is an example of a complex tissue, and is made of two specialized conducting tissues: xylem and phloem. Xylem tissue transports water and nutrients from the roots to different parts of the plant, and includes three different cell types: vessel elements and tracheids (both of which conduct water), and xylem parenchyma. Phloem tissue, which transports organic compounds from the site of photosynthesis to other parts of the plant, consists of four different cell types: sieve cells (which conduct photosynthates), companion cells, phloem parenchyma, and phloem fibers. Unlike xylem conducting cells, phloem conducting cells are alive at maturity. The xylem and phloem always lie adjacent to each other (Figure \(2\)). In stems, the xylem and the phloem form a structure called a vascular bundle; in roots, this is termed the vascular stele or vascular cylinder.
Summary
A vascular plant consists of two organ systems: the shoot system and the root system. The shoot system includes the aboveground vegetative portions (stems and leaves) and reproductive parts (flowers and fruits). The root system supports the plant and is usually underground. A plant is composed of two main types of tissue: meristematic tissue and permanent tissue. Meristematic tissue consists of actively dividing cells found in root and shoot tips. As growth occurs, meristematic tissue differentiates into permanent tissue, which is categorized as either simple or complex. Simple tissues are made up of similar cell types; examples include dermal tissue and ground tissue. Dermal tissue provides the outer covering of the plant. Ground tissue is responsible for photosynthesis; it also supports vascular tissue and may store water and sugars. Complex tissues are made up of different cell types. Vascular tissue, for example, is made up of xylem and phloem cells.
Glossary
apical meristem
meristematic tissue located at the tips of stems and roots; enables a plant to extend in length
dermal tissue
protective plant tissue covering the outermost part of the plant; controls gas exchange
ground tissue
plant tissue involved in photosynthesis; provides support, and stores water and sugars
intercalary meristem
meristematic tissue located at nodes and the bases of leaf blades; found only in monocots
lateral meristem
meristematic tissue that enables a plant to increase in thickness or girth
meristematic tissue
tissue containing cells that constantly divide; contributes to plant growth
meristem
plant region of continuous growth
permanent tissue
plant tissue composed of cells that are no longer actively dividing
root system
belowground portion of the plant that supports the plant and absorbs water and minerals
shoot system
aboveground portion of the plant; consists of non-reproductive plant parts, such as leaves and stems, and reproductive parts, such as flowers and fruits
vascular bundle
strands of stem tissue made up of xylem and phloem
vascular stele
strands of root tissue made up of xylem and phloem
vascular tissue
tissue made up of xylem and phloem that transports food and water throughout the plant | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.1%3A_The_Plant_Body.txt |
Skills to Develop
• Describe the main function and basic structure of stems
• Compare and contrast the roles of dermal tissue, vascular tissue, and ground tissue
• Distinguish between primary growth and secondary growth in stems
• Summarize the origin of annual rings
• List and describe examples of modified stems
Stems are a part of the shoot system of a plant. They may range in length from a few millimeters to hundreds of meters, and also vary in diameter, depending on the plant type. Stems are usually above ground, although the stems of some plants, such as the potato, also grow underground. Stems may be herbaceous (soft) or woody in nature. Their main function is to provide support to the plant, holding leaves, flowers and buds; in some cases, stems also store food for the plant. A stem may be unbranched, like that of a palm tree, or it may be highly branched, like that of a magnolia tree. The stem of the plant connects the roots to the leaves, helping to transport absorbed water and minerals to different parts of the plant. It also helps to transport the products of photosynthesis, namely sugars, from the leaves to the rest of the plant.
Plant stems, whether above or below ground, are characterized by the presence of nodes and internodes (Figure \(1\)). Nodes are points of attachment for leaves, aerial roots, and flowers. The stem region between two nodes is called an internode. The stalk that extends from the stem to the base of the leaf is the petiole. An axillary bud is usually found in the axil—the area between the base of a leaf and the stem—where it can give rise to a branch or a flower. The apex (tip) of the shoot contains the apical meristem within the apical bud.
Stem Anatomy
The stem and other plant organs arise from the ground tissue, and are primarily made up of simple tissues formed from three types of cells: parenchyma, collenchyma, and sclerenchyma cells.
Parenchyma cells are the most common plant cells (Figure \(2\)). They are found in the stem, the root, the inside of the leaf, and the pulp of the fruit. Parenchyma cells are responsible for metabolic functions, such as photosynthesis, and they help repair and heal wounds. Some parenchyma cells also store starch.
Collenchyma cells are elongated cells with unevenly thickened walls (Figure \(3\)). They provide structural support, mainly to the stem and leaves. These cells are alive at maturity and are usually found below the epidermis. The “strings” of a celery stalk are an example of collenchyma cells.
Sclerenchyma cells also provide support to the plant, but unlike collenchyma cells, many of them are dead at maturity. There are two types of sclerenchyma cells: fibers and sclereids. Both types have secondary cell walls that are thickened with deposits of lignin, an organic compound that is a key component of wood. Fibers are long, slender cells; sclereids are smaller-sized. Sclereids give pears their gritty texture. Humans use sclerenchyma fibers to make linen and rope (Figure \(4\)).
Art Connection
Which layers of the stem are made of parenchyma cells?
1. cortex and pith
2. phloem
3. sclerenchyma
4. xylem
Like the rest of the plant, the stem has three tissue systems: dermal, vascular, and ground tissue. Each is distinguished by characteristic cell types that perform specific tasks necessary for the plant’s growth and survival.
Dermal Tissue
The dermal tissue of the stem consists primarily of epidermis, a single layer of cells covering and protecting the underlying tissue. Woody plants have a tough, waterproof outer layer of cork cells commonly known as bark, which further protects the plant from damage. Epidermal cells are the most numerous and least differentiated of the cells in the epidermis. The epidermis of a leaf also contains openings known as stomata, through which the exchange of gases takes place (Figure \(5\)). Two cells, known as guard cells, surround each leaf stoma, controlling its opening and closing and thus regulating the uptake of carbon dioxide and the release of oxygen and water vapor. Trichomes are hair-like structures on the epidermal surface. They help to reduce transpiration (the loss of water by aboveground plant parts), increase solar reflectance, and store compounds that defend the leaves against predation by herbivores.
Vascular Tissue
The xylem and phloem that make up the vascular tissue of the stem are arranged in distinct strands called vascular bundles, which run up and down the length of the stem. When the stem is viewed in cross section, the vascular bundles of dicot stems are arranged in a ring. In plants with stems that live for more than one year, the individual bundles grow together and produce the characteristic growth rings. In monocot stems, the vascular bundles are randomly scattered throughout the ground tissue (Figure \(6\)).
Xylem tissue has three types of cells: xylem parenchyma, tracheids, and vessel elements. The latter two types conduct water and are dead at maturity. Tracheids are xylem cells with thick secondary cell walls that are lignified. Water moves from one tracheid to another through regions on the side walls known as pits, where secondary walls are absent. Vessel elements are xylem cells with thinner walls; they are shorter than tracheids. Each vessel element is connected to the next by means of a perforation plate at the end walls of the element. Water moves through the perforation plates to travel up the plant.
Phloem tissue is composed of sieve-tube cells, companion cells, phloem parenchyma, and phloem fibers. A series of sieve-tube cells (also called sieve-tube elements) are arranged end to end to make up a long sieve tube, which transports organic substances such as sugars and amino acids. The sugars flow from one sieve-tube cell to the next through perforated sieve plates, which are found at the end junctions between two cells. Although still alive at maturity, the nucleus and other cell components of the sieve-tube cells have disintegrated. Companion cells are found alongside the sieve-tube cells, providing them with metabolic support. The companion cells contain more ribosomes and mitochondria than the sieve-tube cells, which lack some cellular organelles.
Ground Tissue
Ground tissue is mostly made up of parenchyma cells, but may also contain collenchyma and sclerenchyma cells that help support the stem. The ground tissue towards the interior of the vascular tissue in a stem or root is known as pith, while the layer of tissue between the vascular tissue and the epidermis is known as the cortex.
Growth in Stems
Growth in plants occurs as the stems and roots lengthen. Some plants, especially those that are woody, also increase in thickness during their life span. The increase in length of the shoot and the root is referred to as primary growth, and is the result of cell division in the shoot apical meristem. Secondary growth is characterized by an increase in thickness or girth of the plant, and is caused by cell division in the lateral meristem. Figure \(7\) shows the areas of primary and secondary growth in a plant. Herbaceous plants mostly undergo primary growth, with hardly any secondary growth or increase in thickness. Secondary growth or “wood” is noticeable in woody plants; it occurs in some dicots, but occurs very rarely in monocots.
Some plant parts, such as stems and roots, continue to grow throughout a plant’s life: a phenomenon called indeterminate growth. Other plant parts, such as leaves and flowers, exhibit determinate growth, which ceases when a plant part reaches a particular size.
Primary Growth
Most primary growth occurs at the apices, or tips, of stems and roots. Primary growth is a result of rapidly dividing cells in the apical meristems at the shoot tip and root tip. Subsequent cell elongation also contributes to primary growth. The growth of shoots and roots during primary growth enables plants to continuously seek water (roots) or sunlight (shoots).
The influence of the apical bud on overall plant growth is known as apical dominance, which diminishes the growth of axillary buds that form along the sides of branches and stems. Most coniferous trees exhibit strong apical dominance, thus producing the typical conical Christmas tree shape. If the apical bud is removed, then the axillary buds will start forming lateral branches. Gardeners make use of this fact when they prune plants by cutting off the tops of branches, thus encouraging the axillary buds to grow out, giving the plant a bushy shape.
Link to Learning
Watch this BBC Nature video showing how time-lapse photography captures plant growth at high speed.
Secondary Growth
The increase in stem thickness that results from secondary growth is due to the activity of the lateral meristems, which are lacking in herbaceous plants. Lateral meristems include the vascular cambium and, in woody plants, the cork cambium (see Figure \(8\)). The vascular cambium is located just outside the primary xylem and to the interior of the primary phloem. The cells of the vascular cambium divide and form secondary xylem (tracheids and vessel elements) to the inside, and secondary phloem (sieve elements and companion cells) to the outside. The thickening of the stem that occurs in secondary growth is due to the formation of secondary phloem and secondary xylem by the vascular cambium, plus the action of cork cambium, which forms the tough outermost layer of the stem. The cells of the secondary xylem contain lignin, which provides hardiness and strength.
In woody plants, cork cambium is the outermost lateral meristem. It produces cork cells (bark) containing a waxy substance known as suberin that can repel water. The bark protects the plant against physical damage and helps reduce water loss. The cork cambium also produces a layer of cells known as phelloderm, which grows inward from the cambium. The cork cambium, cork cells, and phelloderm are collectively termed the periderm. The periderm substitutes for the epidermis in mature plants. In some plants, the periderm has many openings, known as lenticels, which allow the interior cells to exchange gases with the outside atmosphere (Figure \(8\)). This supplies oxygen to the living and metabolically active cells of the cortex, xylem and phloem.
Annual Rings
The activity of the vascular cambium gives rise to annual growth rings. During the spring growing season, cells of the secondary xylem have a large internal diameter and their primary cell walls are not extensively thickened. This is known as early wood, or spring wood. During the fall season, the secondary xylem develops thickened cell walls, forming late wood, or autumn wood, which is denser than early wood. This alternation of early and late wood is due largely to a seasonal decrease in the number of vessel elements and a seasonal increase in the number of tracheids. It results in the formation of an annual ring, which can be seen as a circular ring in the cross section of the stem (Figure \(9\)). An examination of the number of annual rings and their nature (such as their size and cell wall thickness) can reveal the age of the tree and the prevailing climatic conditions during each season.
Stem Modifications
Some plant species have modified stems that are especially suited to a particular habitat and environment (Figure \(10\)). A rhizome is a modified stem that grows horizontally underground and has nodes and internodes. Vertical shoots may arise from the buds on the rhizome of some plants, such as ginger and ferns. Corms are similar to rhizomes, except they are more rounded and fleshy (such as in gladiolus). Corms contain stored food that enables some plants to survive the winter. Stolons are stems that run almost parallel to the ground, or just below the surface, and can give rise to new plants at the nodes. Runners are a type of stolon that runs above the ground and produces new clone plants at nodes at varying intervals: strawberries are an example. Tubers are modified stems that may store starch, as seen in the potato (Solanum sp.). Tubers arise as swollen ends of stolons, and contain many adventitious or unusual buds (familiar to us as the “eyes” on potatoes). A bulb, which functions as an underground storage unit, is a modification of a stem that has the appearance of enlarged fleshy leaves emerging from the stem or surrounding the base of the stem, as seen in the iris.
Link to Learning
Watch botanist Wendy Hodgson, of Desert Botanical Garden in Phoenix, Arizona, explain how agave plants were cultivated for food hundreds of years ago in the Arizona desert in this video: Finding the Roots of an Ancient Crop.
Some aerial modifications of stems are tendrils and thorns (Figure \(11\)). Tendrils are slender, twining strands that enable a plant (like a vine or pumpkin) to seek support by climbing on other surfaces. Thorns are modified branches appearing as sharp outgrowths that protect the plant; common examples include roses, Osage orange and devil’s walking stick.
Summary
The stem of a plant bears the leaves, flowers, and fruits. Stems are characterized by the presence of nodes (the points of attachment for leaves or branches) and internodes (regions between nodes).
Plant organs are made up of simple and complex tissues. The stem has three tissue systems: dermal, vascular, and ground tissue. Dermal tissue is the outer covering of the plant. It contains epidermal cells, stomata, guard cells, and trichomes. Vascular tissue is made up of xylem and phloem tissues and conducts water, minerals, and photosynthetic products. Ground tissue is responsible for photosynthesis and support and is composed of parenchyma, collenchyma, and sclerenchyma cells.
Primary growth occurs at the tips of roots and shoots, causing an increase in length. Woody plants may also exhibit secondary growth, or increase in thickness. In woody plants, especially trees, annual rings may form as growth slows at the end of each season. Some plant species have modified stems that help to store food, propagate new plants, or discourage predators. Rhizomes, corms, stolons, runners, tubers, bulbs, tendrils, and thorns are examples of modified stems.
Art Connections
Figure \(4\): Which layers of the stem are made of parenchyma cells?
1. cortex and pith
2. epidermis
3. sclerenchyma
4. epidermis and cortex.
Answer
A and B. The cortex, pith, and epidermis are made of parenchyma cells.
Glossary
apical bud
bud formed at the tip of the shoot
axillary bud
bud located in the axil: the stem area where the petiole connects to the stem
bark
tough, waterproof, outer epidermal layer of cork cells
bulb
modified underground stem that consists of a large bud surrounded by numerous leaf scales
collenchyma cell
elongated plant cell with unevenly thickened walls; provides structural support to the stem and leaves
companion cell
phloem cell that is connected to sieve-tube cells; has large amounts of ribosomes and mitochondrion
corm
rounded, fleshy underground stem that contains stored food
cortex
ground tissue found between the vascular tissue and the epidermis in a stem or root
epidermis
single layer of cells found in plant dermal tissue; covers and protects underlying tissue
guard cells
paired cells on either side of a stoma that control stomatal opening and thereby regulate the movement of gases and water vapor
internode
region between nodes on the stem
lenticel
opening on the surface of mature woody stems that facilitates gas exchange
node
point along the stem at which leaves, flowers, or aerial roots originate
parenchyma cell
most common type of plant cell; found in the stem, root, leaf, and in fruit pulp; site of photosynthesis and starch storage
periderm
outermost covering of woody stems; consists of the cork cambium, cork cells, and the phelloderm
pith
ground tissue found towards the interior of the vascular tissue in a stem or root
primary growth
growth resulting in an increase in length of the stem and the root; caused by cell division in the shoot or root apical meristem
rhizome
modified underground stem that grows horizontally to the soil surface and has nodes and internodes
runner
stolon that runs above the ground and produces new clone plants at nodes
sclerenchyma cell
plant cell that has thick secondary walls and provides structural support; usually dead at maturity
secondary growth
growth resulting in an increase in thickness or girth; caused by the lateral meristem and cork cambium
sieve-tube cell
phloem cell arranged end to end to form a sieve tube that transports organic substances such as sugars and amino acids
stolon
modified stem that runs parallel to the ground and can give rise to new plants at the nodes
tendril
modified stem consisting of slender, twining strands used for support or climbing
thorn
modified stem branch appearing as a sharp outgrowth that protects the plant
tracheid
xylem cell with thick secondary walls that helps transport water
trichome
hair-like structure on the epidermal surface
tuber
modified underground stem adapted for starch storage; has many adventitious buds
vessel element
xylem cell that is shorter than a tracheid and has thinner walls | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.2%3A_Stems.txt |
Skills to Develop
• Identify the two types of root systems
• Describe the three zones of the root tip and summarize the role of each zone in root growth
• Describe the structure of the root
• List and describe examples of modified roots
The roots of seed plants have three major functions: anchoring the plant to the soil, absorbing water and minerals and transporting them upwards, and storing the products of photosynthesis. Some roots are modified to absorb moisture and exchange gases. Most roots are underground. Some plants, however, also have adventitious roots, which emerge above the ground from the shoot.
Types of Root Systems
Root systems are mainly of two types (Figure \(1\)). Dicots have a tap root system, while monocots have a fibrous root system. A tap root system has a main root that grows down vertically, and from which many smaller lateral roots arise. Dandelions are a good example; their tap roots usually break off when trying to pull these weeds, and they can regrow another shoot from the remaining root). A tap root system penetrates deep into the soil. In contrast, a fibrous root system is located closer to the soil surface, and forms a dense network of roots that also helps prevent soil erosion (lawn grasses are a good example, as are wheat, rice, and corn). Some plants have a combination of tap roots and fibrous roots. Plants that grow in dry areas often have deep root systems, whereas plants growing in areas with abundant water are likely to have shallower root systems.
Root Growth and Anatomy
Root growth begins with seed germination. When the plant embryo emerges from the seed, the radicle of the embryo forms the root system. The tip of the root is protected by the root cap, a structure exclusive to roots and unlike any other plant structure. The root cap is continuously replaced because it gets damaged easily as the root pushes through soil. The root tip can be divided into three zones: a zone of cell division, a zone of elongation, and a zone of maturation and differentiation (Figure \(2\)). The zone of cell division is closest to the root tip; it is made up of the actively dividing cells of the root meristem. The zone of elongation is where the newly formed cells increase in length, thereby lengthening the root. Beginning at the first root hair is the zone of cell maturation where the root cells begin to differentiate into special cell types. All three zones are in the first centimeter or so of the root tip.
The root has an outer layer of cells called the epidermis, which surrounds areas of ground tissue and vascular tissue. The epidermis provides protection and helps in absorption. Root hairs, which are extensions of root epidermal cells, increase the surface area of the root, greatly contributing to the absorption of water and minerals.
Inside the root, the ground tissue forms two regions: the cortex and the pith (Figure \(3\)). Compared to stems, roots have lots of cortex and little pith. Both regions include cells that store photosynthetic products. The cortex is between the epidermis and the vascular tissue, whereas the pith lies between the vascular tissue and the center of the root.
The vascular tissue in the root is arranged in the inner portion of the root, which is called the stele (Figure \(4\)). A layer of cells known as the endodermis separates the stele from the ground tissue in the outer portion of the root. The endodermis is exclusive to roots, and serves as a checkpoint for materials entering the root’s vascular system. A waxy substance called suberin is present on the walls of the endodermal cells. This waxy region, known as the Casparian strip, forces water and solutes to cross the plasma membranes of endodermal cells instead of slipping between the cells. This ensures that only materials required by the root pass through the endodermis, while toxic substances and pathogens are generally excluded. The outermost cell layer of the root’s vascular tissue is the pericycle, an area that can give rise to lateral roots. In dicot roots, the xylem and phloem of the stele are arranged alternately in an X shape, whereas in monocot roots, the vascular tissue is arranged in a ring around the pith.
Root Modifications
Root structures may be modified for specific purposes. For example, some roots are bulbous and store starch. Aerial roots and prop roots are two forms of aboveground roots that provide additional support to anchor the plant. Tap roots, such as carrots, turnips, and beets, are examples of roots that are modified for food storage (Figure \(5\)).
Epiphytic roots enable a plant to grow on another plant. For example, the epiphytic roots of orchids develop a spongy tissue to absorb moisture. The banyan tree (Ficus sp.) begins as an epiphyte, germinating in the branches of a host tree; aerial roots develop from the branches and eventually reach the ground, providing additional support (Figure \(6\)). In screwpine (Pandanus sp.), a palm-like tree that grows in sandy tropical soils, aboveground prop roots develop from the nodes to provide additional support.
Summary
Roots help to anchor a plant, absorb water and minerals, and serve as storage sites for food. Taproots and fibrous roots are the two main types of root systems. In a taproot system, a main root grows vertically downward with a few lateral roots. Fibrous root systems arise at the base of the stem, where a cluster of roots forms a dense network that is shallower than a taproot. The growing root tip is protected by a root cap. The root tip has three main zones: a zone of cell division (cells are actively dividing), a zone of elongation (cells increase in length), and a zone of maturation (cells differentiate to form different kinds of cells). Root vascular tissue conducts water, minerals, and sugars. In some habitats, the roots of certain plants may be modified to form aerial roots or epiphytic roots.
Glossary
adventitious root
aboveground root that arises from a plant part other than the radicle of the plant embryo
Casparian strip
waxy coating that forces water to cross endodermal plasma membranes before entering the vascular cylinder, instead of moving between endodermal cells
endodermis
layer of cells in the root that forms a selective barrier between the ground tissue and the vascular tissue, allowing water and minerals to enter the root while excluding toxins and pathogens
fibrous root system
type of root system in which the roots arise from the base of the stem in a cluster, forming a dense network of roots; found in monocots
pericycle
outer boundary of the stele from which lateral roots can arise
root cap
protective cells covering the tip of the growing root
root hair
hair-like structure that is an extension of epidermal cells; increases the root surface area and aids in absorption of water and minerals
stele
inner portion of the root containing the vascular tissue; surrounded by the endodermis
tap root system
type of root system with a main root that grows vertically with few lateral roots; found in dicots | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.3%3A_Roots.txt |
Skills to Develop
• Identify the parts of a typical leaf
• Describe the internal structure and function of a leaf
• Compare and contrast simple leaves and compound leaves
• List and describe examples of modified leaves
Leaves are the main sites for photosynthesis: the process by which plants synthesize food. Most leaves are usually green, due to the presence of chlorophyll in the leaf cells. However, some leaves may have different colors, caused by other plant pigments that mask the green chlorophyll. The thickness, shape, and size of leaves are adapted to the environment. Each variation helps a plant species maximize its chances of survival in a particular habitat. Usually, the leaves of plants growing in tropical rainforests have larger surface areas than those of plants growing in deserts or very cold conditions, which are likely to have a smaller surface area to minimize water loss.
Structure of a Typical Leaf
Each leaf typically has a leaf blade called the lamina, which is also the widest part of the leaf. Some leaves are attached to the plant stem by a petiole. Leaves that do not have a petiole and are directly attached to the plant stem are called sessile leaves. Small green appendages usually found at the base of the petiole are known as stipules. Most leaves have a midrib, which travels the length of the leaf and branches to each side to produce veins of vascular tissue. The edge of the leaf is called the margin. Figure \(1\) shows the structure of a typical eudicot leaf.
Within each leaf, the vascular tissue forms veins. The arrangement of veins in a leaf is called the venation pattern. Monocots and dicots differ in their patterns of venation (Figure \(2\)). Monocots have parallel venation; the veins run in straight lines across the length of the leaf without converging at a point. In dicots, however, the veins of the leaf have a net-like appearance, forming a pattern known as reticulate venation. One extant plant, the Ginkgo biloba, has dichotomous venation where the veins fork.
Leaf Arrangement
The arrangement of leaves on a stem is known as phyllotaxy. The number and placement of a plant’s leaves will vary depending on the species, with each species exhibiting a characteristic leaf arrangement. Leaves are classified as either alternate, spiral, or opposite. Plants that have only one leaf per node have leaves that are said to be either alternate—meaning the leaves alternate on each side of the stem in a flat plane—or spiral, meaning the leaves are arrayed in a spiral along the stem. In an opposite leaf arrangement, two leaves arise at the same point, with the leaves connecting opposite each other along the branch. If there are three or more leaves connected at a node, the leaf arrangement is classified as whorled.
Leaf Form
Leaves may be simple or compound (Figure \(3\)). In a simple leaf, the blade is either completely undivided—as in the banana leaf—or it has lobes, but the separation does not reach the midrib, as in the maple leaf. In a compound leaf, the leaf blade is completely divided, forming leaflets, as in the locust tree. Each leaflet may have its own stalk, but is attached to the rachis. A palmately compound leaf resembles the palm of a hand, with leaflets radiating outwards from one point Examples include the leaves of poison ivy, the buckeye tree, or the familiar houseplant Schefflera sp. (common name “umbrella plant”). Pinnately compound leaves take their name from their feather-like appearance; the leaflets are arranged along the midrib, as in rose leaves (Rosa sp.), or the leaves of hickory, pecan, ash, or walnut trees.
Leaf Structure and Function
The outermost layer of the leaf is the epidermis; it is present on both sides of the leaf and is called the upper and lower epidermis, respectively. Botanists call the upper side the adaxial surface (or adaxis) and the lower side the abaxial surface (or abaxis). The epidermis helps in the regulation of gas exchange. It contains stomata (Figure \(4\)): openings through which the exchange of gases takes place. Two guard cells surround each stoma, regulating its opening and closing.
The epidermis is usually one cell layer thick; however, in plants that grow in very hot or very cold conditions, the epidermis may be several layers thick to protect against excessive water loss from transpiration. A waxy layer known as the cuticle covers the leaves of all plant species. The cuticle reduces the rate of water loss from the leaf surface. Other leaves may have small hairs (trichomes) on the leaf surface. Trichomes help to deter herbivory by restricting insect movements, or by storing toxic or bad-tasting compounds; they can also reduce the rate of transpiration by blocking air flow across the leaf surface (Figure \(5\)).
Below the epidermis of dicot leaves are layers of cells known as the mesophyll, or “middle leaf.” The mesophyll of most leaves typically contains two arrangements of parenchyma cells: the palisade parenchyma and spongy parenchyma (Figure \(6\)). The palisade parenchyma (also called the palisade mesophyll) has column-shaped, tightly packed cells, and may be present in one, two, or three layers. Below the palisade parenchyma are loosely arranged cells of an irregular shape. These are the cells of the spongy parenchyma (or spongy mesophyll). The air space found between the spongy parenchyma cells allows gaseous exchange between the leaf and the outside atmosphere through the stomata. In aquatic plants, the intercellular spaces in the spongy parenchyma help the leaf float. Both layers of the mesophyll contain many chloroplasts. Guard cells are the only epidermal cells to contain chloroplasts.
Like the stem, the leaf contains vascular bundles composed of xylem and phloem (Figure \(7\)). The xylem consists of tracheids and vessels, which transport water and minerals to the leaves. The phloem transports the photosynthetic products from the leaf to the other parts of the plant. A single vascular bundle, no matter how large or small, always contains both xylem and phloem tissues.
Leaf Adaptations
Coniferous plant species that thrive in cold environments, like spruce, fir, and pine, have leaves that are reduced in size and needle-like in appearance. These needle-like leaves have sunken stomata and a smaller surface area: two attributes that aid in reducing water loss. In hot climates, plants such as cacti have leaves that are reduced to spines, which in combination with their succulent stems, help to conserve water. Many aquatic plants have leaves with wide lamina that can float on the surface of the water, and a thick waxy cuticle on the leaf surface that repels water.
Link to Learning
Watch “The Pale Pitcher Plant” episode of the video series Plants Are Cool, Too, a Botanical Society of America video about a carnivorous plant species found in Louisiana.
Evolution Connection: Plant Adaptations in Resource-Deficient Environments
Roots, stems, and leaves are structured to ensure that a plant can obtain the required sunlight, water, soil nutrients, and oxygen resources. Some remarkable adaptations have evolved to enable plant species to thrive in less than ideal habitats, where one or more of these resources is in short supply.
In tropical rainforests, light is often scarce, since many trees and plants grow close together and block much of the sunlight from reaching the forest floor. Many tropical plant species have exceptionally broad leaves to maximize the capture of sunlight. Other species are epiphytes: plants that grow on other plants that serve as a physical support. Such plants are able to grow high up in the canopy atop the branches of other trees, where sunlight is more plentiful. Epiphytes live on rain and minerals collected in the branches and leaves of the supporting plant. Bromeliads (members of the pineapple family), ferns, and orchids are examples of tropical epiphytes (Figure \(8\)). Many epiphytes have specialized tissues that enable them to efficiently capture and store water.
Some plants have special adaptations that help them to survive in nutrient-poor environments. Carnivorous plants, such as the Venus flytrap and the pitcher plant (Figure \(9\)), grow in bogs where the soil is low in nitrogen. In these plants, leaves are modified to capture insects. The insect-capturing leaves may have evolved to provide these plants with a supplementary source of much-needed nitrogen.
Many swamp plants have adaptations that enable them to thrive in wet areas, where their roots grow submerged underwater. In these aquatic areas, the soil is unstable and little oxygen is available to reach the roots. Trees such as mangroves (Rhizophora sp.) growing in coastal waters produce aboveground roots that help support the tree (Figure \(10\)). Some species of mangroves, as well as cypress trees, have pneumatophores: upward-growing roots containing pores and pockets of tissue specialized for gas exchange. Wild rice is an aquatic plant with large air spaces in the root cortex. The air-filled tissue—called aerenchyma—provides a path for oxygen to diffuse down to the root tips, which are embedded in oxygen-poor bottom sediments.
Link to Learning
Watch Venus Flytraps: Jaws of Death, an extraordinary BBC close-up of the Venus flytrap in action.
Summary
Leaves are the main site of photosynthesis. A typical leaf consists of a lamina (the broad part of the leaf, also called the blade) and a petiole (the stalk that attaches the leaf to a stem). The arrangement of leaves on a stem, known as phyllotaxy, enables maximum exposure to sunlight. Each plant species has a characteristic leaf arrangement and form. The pattern of leaf arrangement may be alternate, opposite, or spiral, while leaf form may be simple or compound. Leaf tissue consists of the epidermis, which forms the outermost cell layer, and mesophyll and vascular tissue, which make up the inner portion of the leaf. In some plant species, leaf form is modified to form structures such as tendrils, spines, bud scales, and needles.
Glossary
compound leaf
leaf in which the leaf blade is subdivided to form leaflets, all attached to the midrib
cuticle
waxy protective layer on the leaf surface
lamina
leaf blade
palmately compound leaf
leaf type with leaflets that emerge from a point, resembling the palm of a hand
petiole
stalk of the leaf
phyllotaxy
arrangement of leaves on a stem
pinnately compound leaf
leaf type with a divided leaf blade consisting of leaflets arranged on both sides of the midrib
sessile
leaf without a petiole that is attached directly to the plant stem
simple leaf
leaf type in which the lamina is completely undivided or merely lobed
stipule
small green structure found on either side of the leaf stalk or petiole
venation
pattern of veins in a leaf; may be parallel (as in monocots), reticulate (as in dicots), or dichotomous (as in Gingko biloba)
whorled
pattern of leaf arrangement in which three or more leaves are connected at a node | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.4%3A_Leaves.txt |
Skills to Develop
• Define water potential and explain how it is influenced by solutes, pressure, gravity, and the matric potential
• Describe how water potential, evapotranspiration, and stomatal regulation influence how water is transported in plants
• Explain how photosynthates are transported in plants
The structure of plant roots, stems, and leaves facilitates the transport of water, nutrients, and photosynthates throughout the plant. The phloem and xylem are the main tissues responsible for this movement. Water potential, evapotranspiration, and stomatal regulation influence how water and nutrients are transported in plants. To understand how these processes work, we must first understand the energetics of water potential.
Water Potential
Plants are phenomenal hydraulic engineers. Using only the basic laws of physics and the simple manipulation of potential energy, plants can move water to the top of a 116-meter-tall tree (Figure $1$a). Plants can also use hydraulics to generate enough force to split rocks and buckle sidewalks (Figure $1$b). Plants achieve this because of water potential.
Water potential is a measure of the potential energy in water. Plant physiologists are not interested in the energy in any one particular aqueous system, but are very interested in water movement between two systems. In practical terms, therefore, water potential is the difference in potential energy between a given water sample and pure water (at atmospheric pressure and ambient temperature). Water potential is denoted by the Greek letter ψ (psi) and is expressed in units of pressure (pressure is a form of energy) called megapascals (MPa). The potential of pure water (Ψwpure H2O) is, by convenience of definition, designated a value of zero (even though pure water contains plenty of potential energy, that energy is ignored). Water potential values for the water in a plant root, stem, or leaf are therefore expressed relative to Ψwpure H2O.
The water potential in plant solutions is influenced by solute concentration, pressure, gravity, and factors called matrix effects. Water potential can be broken down into its individual components using the following equation:
$\psi_\text{system} = \psi_\text{total} = \psi_s + \psi_p + \psi_g + \psi_m \nonumber$
where Ψs, Ψp, Ψg, and Ψm refer to the solute, pressure, gravity, and matric potentials, respectively. “System” can refer to the water potential of the soil water (Ψsoil), root water (Ψroot), stem water (Ψstem), leaf water (Ψleaf) or the water in the atmosphere (Ψatmosphere): whichever aqueous system is under consideration. As the individual components change, they raise or lower the total water potential of a system. When this happens, water moves to equilibrate, moving from the system or compartment with a higher water potential to the system or compartment with a lower water potential. This brings the difference in water potential between the two systems (ΔΨ) back to zero (ΔΨ = 0). Therefore, for water to move through the plant from the soil to the air (a process called transpiration), Ψsoil must be > Ψroot > Ψstem > Ψleaf > Ψatmosphere.
Water only moves in response to ΔΨ, not in response to the individual components. However, because the individual components influence the total Ψsystem, by manipulating the individual components (especially Ψs), a plant can control water movement.
Solute Potential
Solute potential (Ψs), also called osmotic potential, is negative in a plant cell and zero in distilled water. Typical values for cell cytoplasm are –0.5 to –1.0 MPa. Solutes reduce water potential (resulting in a negative Ψw) by consuming some of the potential energy available in the water. Solute molecules can dissolve in water because water molecules can bind to them via hydrogen bonds; a hydrophobic molecule like oil, which cannot bind to water, cannot go into solution. The energy in the hydrogen bonds between solute molecules and water is no longer available to do work in the system because it is tied up in the bond. In other words, the amount of available potential energy is reduced when solutes are added to an aqueous system. Thus, Ψs decreases with increasing solute concentration. Because Ψs is one of the four components of Ψsystem or Ψtotal, a decrease in Ψs will cause a decrease in Ψtotal. The internal water potential of a plant cell is more negative than pure water because of the cytoplasm’s high solute content (Figure $2$). Because of this difference in water potential water will move from the soil into a plant’s root cells via the process of osmosis. This is why solute potential is sometimes called osmotic potential.
Plant cells can metabolically manipulate Ψs (and by extension, Ψtotal) by adding or removing solute molecules. Therefore, plants have control over Ψtotal via their ability to exert metabolic control over Ψs.
Art Connection
Positive water potential is placed on the left side of the tube by increasing Ψp such that the water level rises on the right side. Could you equalize the water level on each side of the tube by adding solute, and if so, how?
Pressure Potential
Pressure potential (Ψp), also called turgor potential, may be positive or negative (Figure $3$). Because pressure is an expression of energy, the higher the pressure, the more potential energy in a system, and vice versa. Therefore, a positive Ψp (compression) increases Ψtotal, and a negative Ψp (tension) decreases Ψtotal. Positive pressure inside cells is contained by the cell wall, producing turgor pressure. Pressure potentials are typically around 0.6–0.8 MPa, but can reach as high as 1.5 MPa in a well-watered plant. A Ψp of 1.5 MPa equates to 210 pounds per square inch (1.5 MPa x 140 lb in-2 MPa-1 = 210 lb/in-2). As a comparison, most automobile tires are kept at a pressure of 30–34 psi. An example of the effect of turgor pressure is the wilting of leaves and their restoration after the plant has been watered (Figure $3$). Water is lost from the leaves via transpiration (approaching Ψp = 0 MPa at the wilting point) and restored by uptake via the roots.
A plant can manipulate Ψp via its ability to manipulate Ψs and by the process of osmosis. If a plant cell increases the cytoplasmic solute concentration, Ψs will decline, Ψtotal will decline, the ΔΨ between the cell and the surrounding tissue will decline, water will move into the cell by osmosis, and Ψp will increase. Ψp is also under indirect plant control via the opening and closing of stomata. Stomatal openings allow water to evaporate from the leaf, reducing Ψp and Ψtotal of the leaf and increasing ii between the water in the leaf and the petiole, thereby allowing water to flow from the petiole into the leaf.
Gravity Potential
Gravity potential (Ψg) is always negative to zero in a plant with no height. It always removes or consumes potential energy from the system. The force of gravity pulls water downwards to the soil, reducing the total amount of potential energy in the water in the plant (Ψtotal). The taller the plant, the taller the water column, and the more influential Ψg becomes. On a cellular scale and in short plants, this effect is negligible and easily ignored. However, over the height of a tall tree like a giant coastal redwood, the gravitational pull of –0.1 MPa m-1 is equivalent to an extra 1 MPa of resistance that must be overcome for water to reach the leaves of the tallest trees. Plants are unable to manipulate Ψg.
Matric Potential
Matric potential (Ψm) is always negative to zero. In a dry system, it can be as low as –2 MPa in a dry seed, and it is zero in a water-saturated system. The binding of water to a matrix always removes or consumes potential energy from the system. Ψm is similar to solute potential because it involves tying up the energy in an aqueous system by forming hydrogen bonds between the water and some other component. However, in solute potential, the other components are soluble, hydrophilic solute molecules, whereas in Ψm, the other components are insoluble, hydrophilic molecules of the plant cell wall. Every plant cell has a cellulosic cell wall and the cellulose in the cell walls is hydrophilic, producing a matrix for adhesion of water: hence the name matric potential. Ψm is very large (negative) in dry tissues such as seeds or drought-affected soils. However, it quickly goes to zero as the seed takes up water or the soil hydrates. Ψm cannot be manipulated by the plant and is typically ignored in well-watered roots, stems, and leaves.
Movement of Water and Minerals in the Xylem
Solutes, pressure, gravity, and matric potential are all important for the transport of water in plants. Water moves from an area of higher total water potential (higher Gibbs free energy) to an area of lower total water potential. Gibbs free energy is the energy associated with a chemical reaction that can be used to do work. This is expressed as ΔΨ.
Transpiration is the loss of water from the plant through evaporation at the leaf surface. It is the main driver of water movement in the xylem. Transpiration is caused by the evaporation of water at the leaf–atmosphere interface; it creates negative pressure (tension) equivalent to –2 MPa at the leaf surface. This value varies greatly depending on the vapor pressure deficit, which can be negligible at high relative humidity (RH) and substantial at low RH. Water from the roots is pulled up by this tension. At night, when stomata shut and transpiration stops, the water is held in the stem and leaf by the adhesion of water to the cell walls of the xylem vessels and tracheids, and the cohesion of water molecules to each other. This is called the cohesion–tension theory of sap ascent.
Inside the leaf at the cellular level, water on the surface of mesophyll cells saturates the cellulose microfibrils of the primary cell wall. The leaf contains many large intercellular air spaces for the exchange of oxygen for carbon dioxide, which is required for photosynthesis. The wet cell wall is exposed to this leaf internal air space, and the water on the surface of the cells evaporates into the air spaces, decreasing the thin film on the surface of the mesophyll cells. This decrease creates a greater tension on the water in the mesophyll cells (Figure $4$), thereby increasing the pull on the water in the xylem vessels. The xylem vessels and tracheids are structurally adapted to cope with large changes in pressure. Rings in the vessels maintain their tubular shape, much like the rings on a vacuum cleaner hose keep the hose open while it is under pressure. Small perforations between vessel elements reduce the number and size of gas bubbles that can form via a process called cavitation. The formation of gas bubbles in xylem interrupts the continuous stream of water from the base to the top of the plant, causing a break termed an embolism in the flow of xylem sap. The taller the tree, the greater the tension forces needed to pull water, and the more cavitation events. In larger trees, the resulting embolisms can plug xylem vessels, making them non-functional.
Art Connection
Which of the following statements is false?
1. Negative water potential draws water into the root hairs. Cohesion and adhesion draw water up the xylem. Transpiration draws water from the leaf.
2. Negative water potential draws water into the root hairs. Cohesion and adhesion draw water up the phloem. Transpiration draws water from the leaf.
3. Water potential decreases from the roots to the top of the plant.
4. Water enters the plants through root hairs and exits through stoma.
Transpiration—the loss of water vapor to the atmosphere through stomata—is a passive process, meaning that metabolic energy in the form of ATP is not required for water movement. The energy driving transpiration is the difference in energy between the water in the soil and the water in the atmosphere. However, transpiration is tightly controlled.
Control of Transpiration
The atmosphere to which the leaf is exposed drives transpiration, but also causes massive water loss from the plant. Up to 90 percent of the water taken up by roots may be lost through transpiration.
Leaves are covered by a waxy cuticle on the outer surface that prevents the loss of water. Regulation of transpiration, therefore, is achieved primarily through the opening and closing of stomata on the leaf surface. Stomata are surrounded by two specialized cells called guard cells, which open and close in response to environmental cues such as light intensity and quality, leaf water status, and carbon dioxide concentrations. Stomata must open to allow air containing carbon dioxide and oxygen to diffuse into the leaf for photosynthesis and respiration. When stomata are open, however, water vapor is lost to the external environment, increasing the rate of transpiration. Therefore, plants must maintain a balance between efficient photosynthesis and water loss.
Plants have evolved over time to adapt to their local environment and reduce transpiration(Figure $5$). Desert plant (xerophytes) and plants that grow on other plants (epiphytes) have limited access to water. Such plants usually have a much thicker waxy cuticle than those growing in more moderate, well-watered environments (mesophytes). Aquatic plants (hydrophytes) also have their own set of anatomical and morphological leaf adaptations.
Xerophytes and epiphytes often have a thick covering of trichomes or of stomata that are sunken below the leaf’s surface. Trichomes are specialized hair-like epidermal cells that secrete oils and substances. These adaptations impede air flow across the stomatal pore and reduce transpiration. Multiple epidermal layers are also commonly found in these types of plants.
Transportation of Photosynthates in the Phloem
Plants need an energy source to grow. In seeds and bulbs, food is stored in polymers (such as starch) that are converted by metabolic processes into sucrose for newly developing plants. Once green shoots and leaves are growing, plants are able to produce their own food by photosynthesizing. The products of photosynthesis are called photosynthates, which are usually in the form of simple sugars such as sucrose.
Structures that produce photosynthates for the growing plant are referred to as sources. Sugars produced in sources, such as leaves, need to be delivered to growing parts of the plant via the phloem in a process called translocation. The points of sugar delivery, such as roots, young shoots, and developing seeds, are called sinks. Seeds, tubers, and bulbs can be either a source or a sink, depending on the plant’s stage of development and the season.
The products from the source are usually translocated to the nearest sink through the phloem. For example, the highest leaves will send photosynthates upward to the growing shoot tip, whereas lower leaves will direct photosynthates downward to the roots. Intermediate leaves will send products in both directions, unlike the flow in the xylem, which is always unidirectional (soil to leaf to atmosphere). The pattern of photosynthate flow changes as the plant grows and develops. Photosynthates are directed primarily to the roots early on, to shoots and leaves during vegetative growth, and to seeds and fruits during reproductive development. They are also directed to tubers for storage.
Translocation: Transport from Source to Sink
Photosynthates, such as sucrose, are produced in the mesophyll cells of photosynthesizing leaves. From there they are translocated through the phloem to where they are used or stored. Mesophyll cells are connected by cytoplasmic channels called plasmodesmata. Photosynthates move through these channels to reach phloem sieve-tube elements (STEs) in the vascular bundles. From the mesophyll cells, the photosynthates are loaded into the phloem STEs. The sucrose is actively transported against its concentration gradient (a process requiring ATP) into the phloem cells using the electrochemical potential of the proton gradient. This is coupled to the uptake of sucrose with a carrier protein called the sucrose-H+ symporter.
Phloem STEs have reduced cytoplasmic contents, and are connected by a sieve plate with pores that allow for pressure-driven bulk flow, or translocation, of phloem sap. Companion cells are associated with STEs. They assist with metabolic activities and produce energy for the STEs (Figure $6$).
Once in the phloem, the photosynthates are translocated to the closest sink. Phloem sap is an aqueous solution that contains up to 30 percent sugar, minerals, amino acids, and plant growth regulators. The high percentage of sugar decreases Ψs, which decreases the total water potential and causes water to move by osmosis from the adjacent xylem into the phloem tubes, thereby increasing pressure. This increase in total water potential causes the bulk flow of phloem from source to sink (Figure $7$). Sucrose concentration in the sink cells is lower than in the phloem STEs because the sink sucrose has been metabolized for growth, or converted to starch for storage or other polymers, such as cellulose, for structural integrity. Unloading at the sink end of the phloem tube occurs by either diffusion or active transport of sucrose molecules from an area of high concentration to one of low concentration. Water diffuses from the phloem by osmosis and is then transpired or recycled via the xylem back into the phloem sap.
Summary
Water potential (Ψ) is a measure of the difference in potential energy between a water sample and pure water. The water potential in plant solutions is influenced by solute concentration, pressure, gravity, and matric potential. Water potential and transpiration influence how water is transported through the xylem in plants. These processes are regulated by stomatal opening and closing. Photosynthates (mainly sucrose) move from sources to sinks through the plant’s phloem. Sucrose is actively loaded into the sieve-tube elements of the phloem. The increased solute concentration causes water to move by osmosis from the xylem into the phloem. The positive pressure that is produced pushes water and solutes down the pressure gradient. The sucrose is unloaded into the sink, and the water returns to the xylem vessels.
Art Connections
Figure $2$: Positive water potential is placed on the left side of the tube by increasing Ψp such that the water level rises on the right side. Could you equalize the water level on each side of the tube by adding solute, and if so, how?
Answer
Yes, you can equalize the water level by adding the solute to the left side of the tube such that water moves toward the left until the water levels are equal.
Figure $4$: Which of the following statements is false?
1. Negative water potential draws water into the root hairs. Cohesion and adhesion draw water up the xylem. Transpiration draws water from the leaf.
2. Negative water potential draws water into the root hairs. Cohesion and adhesion draw water up the phloem. Transpiration draws water from the leaf.
3. Water potential decreases from the roots to the top of the plant.
4. Water enters the plants through root hairs and exits through stoma.
Answer
B.
Glossary
cuticle
waxy covering on the outside of the leaf and stem that prevents the loss of water
megapascal (MPa)
pressure units that measure water potential
sink
growing parts of a plant, such as roots and young leaves, which require photosynthate
source
organ that produces photosynthate for a plant
translocation
mass transport of photosynthates from source to sink in vascular plants
transpiration
loss of water vapor to the atmosphere through stomata
water potential (Ψw)
the potential energy of a water solution per unit volume in relation to pure water at atmospheric pressure and ambient temperature | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.5%3A_Transport_of_Water_and_Solutes_in_Plants.txt |
Skills to Develop
• Describe how red and blue light affect plant growth and metabolic activities
• Discuss gravitropism
• Understand how hormones affect plant growth and development
• Describe thigmotropism, thigmonastism, and thigmogenesis
• Explain how plants defend themselves from predators and respond to wounds
Animals can respond to environmental factors by moving to a new location. Plants, however, are rooted in place and must respond to the surrounding environmental factors. Plants have sophisticated systems to detect and respond to light, gravity, temperature, and physical touch. Receptors sense environmental factors and relay the information to effector systems—often through intermediate chemical messengers—to bring about plant responses.
Plant Responses to Light
Plants have a number of sophisticated uses for light that go far beyond their ability to photosynthesize low-molecular-weight sugars using only carbon dioxide, light, and water. Photomorphogenesis is the growth and development of plants in response to light. It allows plants to optimize their use of light and space. Photoperiodism is the ability to use light to track time. Plants can tell the time of day and time of year by sensing and using various wavelengths of sunlight. Phototropism is a directional response that allows plants to grow towards, or even away from, light.
The sensing of light in the environment is important to plants; it can be crucial for competition and survival. The response of plants to light is mediated by different photoreceptors, which are comprised of a protein covalently bonded to a light-absorbing pigment called a chromophore. Together, the two are called a chromoprotein.
The red/far-red and violet-blue regions of the visible light spectrum trigger structural development in plants. Sensory photoreceptors absorb light in these particular regions of the visible light spectrum because of the quality of light available in the daylight spectrum. In terrestrial habitats, light absorption by chlorophylls peaks in the blue and red regions of the spectrum. As light filters through the canopy and the blue and red wavelengths are absorbed, the spectrum shifts to the far-red end, shifting the plant community to those plants better adapted to respond to far-red light. Blue-light receptors allow plants to gauge the direction and abundance of sunlight, which is rich in blue–green emissions. Water absorbs red light, which makes the detection of blue light essential for algae and aquatic plants.
The Phytochrome System and the Red/Far-Red Response
The phytochromes are a family of chromoproteins with a linear tetrapyrrole chromophore, similar to the ringed tetrapyrrole light-absorbing head group of chlorophyll. Phytochromes have two photo-interconvertible forms: Pr and Pfr. Pr absorbs red light (~667 nm) and is immediately converted to Pfr. Pfr absorbs far-red light (~730 nm) and is quickly converted back to Pr. Absorption of red or far-red light causes a massive change to the shape of the chromophore, altering the conformation and activity of the phytochrome protein to which it is bound. Pfr is the physiologically active form of the protein; therefore, exposure to red light yields physiological activity. Exposure to far-red light inhibits phytochrome activity. Together, the two forms represent the phytochrome system (Figure \(1\)).
The phytochrome system acts as a biological light switch. It monitors the level, intensity, duration, and color of environmental light. The effect of red light is reversible by immediately shining far-red light on the sample, which converts the chromoprotein to the inactive Pr form. Additionally, Pfr can slowly revert to Pr in the dark, or break down over time. In all instances, the physiological response induced by red light is reversed. The active form of phytochrome (Pfr) can directly activate other molecules in the cytoplasm, or it can be trafficked to the nucleus, where it directly activates or represses specific gene expression.
Once the phytochrome system evolved, plants adapted it to serve a variety of needs. Unfiltered, full sunlight contains much more red light than far-red light. Because chlorophyll absorbs strongly in the red region of the visible spectrum, but not in the far-red region, any plant in the shade of another plant on the forest floor will be exposed to red-depleted, far-red-enriched light. The preponderance of far-red light converts phytochrome in the shaded leaves to the Pr (inactive) form, slowing growth. The nearest non-shaded (or even less-shaded) areas on the forest floor have more red light; leaves exposed to these areas sense the red light, which activates the Pfr form and induces growth. In short, plant shoots use the phytochrome system to grow away from shade and towards light. Because competition for light is so fierce in a dense plant community, the evolutionary advantages of the phytochrome system are obvious.
In seeds, the phytochrome system is not used to determine direction and quality of light (shaded versus unshaded). Instead, is it used merely to determine if there is any light at all. This is especially important in species with very small seeds, such as lettuce. Because of their size, lettuce seeds have few food reserves. Their seedlings cannot grow for long before they run out of fuel. If they germinated even a centimeter under the soil surface, the seedling would never make it into the sunlight and would die. In the dark, phytochrome is in the Pr (inactive form) and the seed will not germinate; it will only germinate if exposed to light at the surface of the soil. Upon exposure to light, Pr is converted to Pfr and germination proceeds.
Plants also use the phytochrome system to sense the change of season. Photoperiodism is a biological response to the timing and duration of day and night. It controls flowering, setting of winter buds, and vegetative growth. Detection of seasonal changes is crucial to plant survival. Although temperature and light intensity influence plant growth, they are not reliable indicators of season because they may vary from one year to the next. Day length is a better indicator of the time of year.
As stated above, unfiltered sunlight is rich in red light but deficient in far-red light. Therefore, at dawn, all the phytochrome molecules in a leaf quickly convert to the active Pfr form, and remain in that form until sunset. In the dark, the Pfr form takes hours to slowly revert back to the Pr form. If the night is long (as in winter), all of the Pfr form reverts. If the night is short (as in summer), a considerable amount of Pfr may remain at sunrise. By sensing the Pr/Pfr ratio at dawn, a plant can determine the length of the day/night cycle. In addition, leaves retain that information for several days, allowing a comparison between the length of the previous night and the preceding several nights. Shorter nights indicate springtime to the plant; when the nights become longer, autumn is approaching. This information, along with sensing temperature and water availability, allows plants to determine the time of the year and adjust their physiology accordingly. Short-day (long-night) plants use this information to flower in the late summer and early fall, when nights exceed a critical length (often eight or fewer hours). Long-day (short-night) plants flower during the spring, when darkness is less than a critical length (often eight to 15 hours). Not all plants use the phytochrome system in this way. Flowering in day-neutral plants is not regulated by daylength.
Career Connection: Horticulturalist
The word “horticulturist” comes from the Latin words for garden (hortus) and culture (cultura). This career has been revolutionized by progress made in the understanding of plant responses to environmental stimuli. Growers of crops, fruit, vegetables, and flowers were previously constrained by having to time their sowing and harvesting according to the season. Now, horticulturists can manipulate plants to increase leaf, flower, or fruit production by understanding how environmental factors affect plant growth and development.
Greenhouse management is an essential component of a horticulturist’s education. To lengthen the night, plants are covered with a blackout shade cloth. Long-day plants are irradiated with red light in winter to promote early flowering. For example, fluorescent (cool white) light high in blue wavelengths encourages leafy growth and is excellent for starting seedlings. Incandescent lamps (standard light bulbs) are rich in red light, and promote flowering in some plants. The timing of fruit ripening can be increased or delayed by applying plant hormones. Recently, considerable progress has been made in the development of plant breeds that are suited to different climates and resistant to pests and transportation damage. Both crop yield and quality have increased as a result of practical applications of the knowledge of plant responses to external stimuli and hormones.
Horticulturists find employment in private and governmental laboratories, greenhouses, botanical gardens, and in the production or research fields. They improve crops by applying their knowledge of genetics and plant physiology. To prepare for a horticulture career, students take classes in botany, plant physiology, plant pathology, landscape design, and plant breeding. To complement these traditional courses, horticulture majors add studies in economics, business, computer science, and communications.
The Blue Light Responses
Phototropism—the directional bending of a plant toward or away from a light source—is a response to blue wavelengths of light. Positive phototropism is growth towards a light source (Figure \(2\)), while negative phototropism (also called skototropism) is growth away from light.
The aptly-named phototropins are protein-based receptors responsible for mediating the phototropic response. Like all plant photoreceptors, phototropins consist of a protein portion and a light-absorbing portion, called the chromophore. In phototropins, the chromophore is a covalently-bound molecule of flavin; hence, phototropins belong to a class of proteins called flavoproteins.
Other responses under the control of phototropins are leaf opening and closing, chloroplast movement, and the opening of stomata. However, of all responses controlled by phototropins, phototropism has been studied the longest and is the best understood.
In their 1880 treatise The Power of Movements in Plants, Charles Darwin and his son Francis first described phototropism as the bending of seedlings toward light. Darwin observed that light was perceived by the tip of the plant (the apical meristem), but that the response (bending) took place in a different part of the plant. They concluded that the signal had to travel from the apical meristem to the base of the plant.
In 1913, Peter Boysen-Jensen demonstrated that a chemical signal produced in the plant tip was responsible for the bending at the base. He cut off the tip of a seedling, covered the cut section with a layer of gelatin, and then replaced the tip. The seedling bent toward the light when illuminated. However, when impermeable mica flakes were inserted between the tip and the cut base, the seedling did not bend. A refinement of the experiment showed that the signal traveled on the shaded side of the seedling. When the mica plate was inserted on the illuminated side, the plant did bend towards the light. Therefore, the chemical signal was a growth stimulant because the phototropic response involved faster cell elongation on the shaded side than on the illuminated side. We now know that as light passes through a plant stem, it is diffracted and generates phototropin activation across the stem. Most activation occurs on the lit side, causing the plant hormone indole acetic acid (IAA) to accumulate on the shaded side. Stem cells elongate under influence of IAA.
Cryptochromes are another class of blue-light absorbing photoreceptors that also contain a flavin-based chromophore. Cryptochromes set the plants 24-hour activity cycle, also know as its circadian rhythem, using blue light cues. There is some evidence that cryptochromes work together with phototropins to mediate the phototropic response.
Link to Learning
Use the navigation menu in the left panel of this website to view images of plants in motion.
Plant Responses to Gravity
Whether or not they germinate in the light or in total darkness, shoots usually sprout up from the ground, and roots grow downward into the ground. A plant laid on its side in the dark will send shoots upward when given enough time. Gravitropism ensures that roots grow into the soil and that shoots grow toward sunlight. Growth of the shoot apical tip upward is called negative gravitropism, whereas growth of the roots downward is called positive gravitropism.
Amyloplasts (also known as statoliths) are specialized plastids that contain starch granules and settle downward in response to gravity. Amyloplasts are found in shoots and in specialized cells of the root cap. When a plant is tilted, the statoliths drop to the new bottom cell wall. A few hours later, the shoot or root will show growth in the new vertical direction.
The mechanism that mediates gravitropism is reasonably well understood. When amyloplasts settle to the bottom of the gravity-sensing cells in the root or shoot, they physically contact the endoplasmic reticulum (ER), causing the release of calcium ions from inside the ER. This calcium signaling in the cells causes polar transport of the plant hormone IAA to the bottom of the cell. In roots, a high concentration of IAA inhibits cell elongation. The effect slows growth on the lower side of the root, while cells develop normally on the upper side. IAA has the opposite effect in shoots, where a higher concentration at the lower side of the shoot stimulates cell expansion, causing the shoot to grow up. After the shoot or root begin to grow vertically, the amyloplasts return to their normal position. Other hypotheses—involving the entire cell in the gravitropism effect—have been proposed to explain why some mutants that lack amyloplasts may still exhibit a weak gravitropic response.
Growth Responses
A plant’s sensory response to external stimuli relies on chemical messengers (hormones). Plant hormones affect all aspects of plant life, from flowering to fruit setting and maturation, and from phototropism to leaf fall. Potentially every cell in a plant can produce plant hormones. They can act in their cell of origin or be transported to other portions of the plant body, with many plant responses involving the synergistic or antagonistic interaction of two or more hormones. In contrast, animal hormones are produced in specific glands and transported to a distant site for action, and they act alone.
Plant hormones are a group of unrelated chemical substances that affect plant morphogenesis. Five major plant hormones are traditionally described: auxins (particularly IAA), cytokinins, gibberellins, ethylene, and abscisic acid. In addition, other nutrients and environmental conditions can be characterized as growth factors.
Auxins
The term auxin is derived from the Greek word auxein, which means "to grow." Auxins are the main hormones responsible for cell elongation in phototropism and gravitropism. They also control the differentiation of meristem into vascular tissue, and promote leaf development and arrangement. While many synthetic auxins are used as herbicides, IAA is the only naturally occurring auxin that shows physiological activity. Apical dominance—the inhibition of lateral bud formation—is triggered by auxins produced in the apical meristem. Flowering, fruit setting and ripening, and inhibition of abscission (leaf falling) are other plant responses under the direct or indirect control of auxins. Auxins also act as a relay for the effects of the blue light and red/far-red responses.
Commercial use of auxins is widespread in plant nurseries and for crop production. IAA is used as a rooting hormone to promote growth of adventitious roots on cuttings and detached leaves. Applying synthetic auxins to tomato plants in greenhouses promotes normal fruit development. Outdoor application of auxin promotes synchronization of fruit setting and dropping to coordinate the harvesting season. Fruits such as seedless cucumbers can be induced to set fruit by treating unfertilized plant flowers with auxins.
Cytokinins
The effect of cytokinins was first reported when it was found that adding the liquid endosperm of coconuts to developing plant embryos in culture stimulated their growth. The stimulating growth factor was found to be cytokinin, a hormone that promotes cytokinesis (cell division). Almost 200 naturally occurring or synthetic cytokinins are known to date. Cytokinins are most abundant in growing tissues, such as roots, embryos, and fruits, where cell division is occurring. Cytokinins are known to delay senescence in leaf tissues, promote mitosis, and stimulate differentiation of the meristem in shoots and roots. Many effects on plant development are under the influence of cytokinins, either in conjunction with auxin or another hormone. For example, apical dominance seems to result from a balance between auxins that inhibit lateral buds, and cytokinins that promote bushier growth.
Gibberellins
Gibberellins (GAs) are a group of about 125 closely related plant hormones that stimulate shoot elongation, seed germination, and fruit and flower maturation. GAs are synthesized in the root and stem apical meristems, young leaves, and seed embryos. In urban areas, GA antagonists are sometimes applied to trees under power lines to control growth and reduce the frequency of pruning.
GAs break dormancy (a state of inhibited growth and development) in the seeds of plants that require exposure to cold or light to germinate. Abscisic acid is a strong antagonist of GA action. Other effects of GAs include gender expression, seedless fruit development, and the delay of senescence in leaves and fruit. Seedless grapes are obtained through standard breeding methods and contain inconspicuous seeds that fail to develop. Because GAs are produced by the seeds, and because fruit development and stem elongation are under GA control, these varieties of grapes would normally produce small fruit in compact clusters. Maturing grapes are routinely treated with GA to promote larger fruit size, as well as looser bunches (longer stems), which reduces the instance of mildew infection (Figure \(3\)).
Abscisic Acid
The plant hormone abscisic acid (ABA) was first discovered as the agent that causes the abscission or dropping of cotton bolls. However, more recent studies indicate that ABA plays only a minor role in the abscission process. ABA accumulates as a response to stressful environmental conditions, such as dehydration, cold temperatures, or shortened day lengths. Its activity counters many of the growth-promoting effects of GAs and auxins. ABA inhibits stem elongation and induces dormancy in lateral buds.
ABA induces dormancy in seeds by blocking germination and promoting the synthesis of storage proteins. Plants adapted to temperate climates require a long period of cold temperature before seeds germinate. This mechanism protects young plants from sprouting too early during unseasonably warm weather in winter. As the hormone gradually breaks down over winter, the seed is released from dormancy and germinates when conditions are favorable in spring. Another effect of ABA is to promote the development of winter buds; it mediates the conversion of the apical meristem into a dormant bud. Low soil moisture causes an increase in ABA, which causes stomata to close, reducing water loss in winter buds.
Ethylene
Ethylene is associated with fruit ripening, flower wilting, and leaf fall. Ethylene is unusual because it is a volatile gas (C2H4). Hundreds of years ago, when gas street lamps were installed in city streets, trees that grew close to lamp posts developed twisted, thickened trunks and shed their leaves earlier than expected. These effects were caused by ethylene volatilizing from the lamps.
Aging tissues (especially senescing leaves) and nodes of stems produce ethylene. The best-known effect of the hormone, however, is the promotion of fruit ripening. Ethylene stimulates the conversion of starch and acids to sugars. Some people store unripe fruit, such as avocadoes, in a sealed paper bag to accelerate ripening; the gas released by the first fruit to mature will speed up the maturation of the remaining fruit. Ethylene also triggers leaf and fruit abscission, flower fading and dropping, and promotes germination in some cereals and sprouting of bulbs and potatoes.
Ethylene is widely used in agriculture. Commercial fruit growers control the timing of fruit ripening with application of the gas. Horticulturalists inhibit leaf dropping in ornamental plants by removing ethylene from greenhouses using fans and ventilation.
Nontraditional Hormones
Recent research has discovered a number of compounds that also influence plant development. Their roles are less understood than the effects of the major hormones described so far.
Jasmonates play a major role in defense responses to herbivory. Their levels increase when a plant is wounded by a predator, resulting in an increase in toxic secondary metabolites. They contribute to the production of volatile compounds that attract natural enemies of predators. For example, chewing of tomato plants by caterpillars leads to an increase in jasmonic acid levels, which in turn triggers the release of volatile compounds that attract predators of the pest.
Oligosaccharins also play a role in plant defense against bacterial and fungal infections. They act locally at the site of injury, and can also be transported to other tissues. Strigolactones promote seed germination in some species and inhibit lateral apical development in the absence of auxins. Strigolactones also play a role in the establishment of mycorrhizae, a mutualistic association of plant roots and fungi. Brassinosteroids are important to many developmental and physiological processes. Signals between these compounds and other hormones, notably auxin and GAs, amplifies their physiological effect. Apical dominance, seed germination, gravitropism, and resistance to freezing are all positively influenced by hormones. Root growth and fruit dropping are inhibited by steroids.
Plant Responses to Wind and Touch
The shoot of a pea plant winds around a trellis, while a tree grows on an angle in response to strong prevailing winds. These are examples of how plants respond to touch or wind.
The movement of a plant subjected to constant directional pressure is called thigmotropism, from the Greek words thigma meaning “touch,” and tropism implying “direction.” Tendrils are one example of this. The meristematic region of tendrils is very touch sensitive; light touch will evoke a quick coiling response. Cells in contact with a support surface contract, whereas cells on the opposite side of the support expand. Application of jasmonic acid is sufficient to trigger tendril coiling without a mechanical stimulus.
A thigmonastic response is a touch response independent of the direction of stimulus. In the Venus flytrap, two modified leaves are joined at a hinge and lined with thin fork-like tines along the outer edges. Tiny hairs are located inside the trap. When an insect brushes against these trigger hairs, touching two or more of them in succession, the leaves close quickly, trapping the prey. Glands on the leaf surface secrete enzymes that slowly digest the insect. The released nutrients are absorbed by the leaves, which reopen for the next meal.
Thigmomorphogenesis is a slow developmental change in the shape of a plant subjected to continuous mechanical stress. When trees bend in the wind, for example, growth is usually stunted and the trunk thickens. Strengthening tissue, especially xylem, is produced to add stiffness to resist the wind’s force. Researchers hypothesize that mechanical strain induces growth and differentiation to strengthen the tissues. Ethylene and jasmonate are likely involved in thigmomorphogenesis.
Link to Learning
Use the menu at the left to navigate to three short movies: a Venus fly trap capturing prey, the progressive closing of sensitive plant leaflets, and the twining of tendrils.
Defense Responses against Herbivores and Pathogens
Plants face two types of enemies: herbivores and pathogens. Herbivores both large and small use plants as food, and actively chew them. Pathogens are agents of disease. These infectious microorganisms, such as fungi, bacteria, and nematodes, live off of the plant and damage its tissues. Plants have developed a variety of strategies to discourage or kill attackers.
The first line of defense in plants is an intact and impenetrable barrier. Bark and the waxy cuticle can protect against predators. Other adaptations against herbivory include thorns, which are modified branches, and spines, which are modified leaves. They discourage animals by causing physical damage and inducing rashes and allergic reactions. A plant’s exterior protection can be compromised by mechanical damage, which may provide an entry point for pathogens. If the first line of defense is breached, the plant must resort to a different set of defense mechanisms, such as toxins and enzymes.
Secondary metabolites are compounds that are not directly derived from photosynthesis and are not necessary for respiration or plant growth and development. Many metabolites are toxic, and can even be lethal to animals that ingest them. Some metabolites are alkaloids, which discourage predators with noxious odors (such as the volatile oils of mint and sage) or repellent tastes (like the bitterness of quinine). Other alkaloids affect herbivores by causing either excessive stimulation (caffeine is one example) or the lethargy associated with opioids. Some compounds become toxic after ingestion; for instance, glycol cyanide in the cassava root releases cyanide only upon ingestion by the herbivore.
Mechanical wounding and predator attacks activate defense and protection mechanisms both in the damaged tissue and at sites farther from the injury location. Some defense reactions occur within minutes: others over several hours. The infected and surrounding cells may die, thereby stopping the spread of infection.
Long-distance signaling elicits a systemic response aimed at deterring the predator. As tissue is damaged, jasmonates may promote the synthesis of compounds that are toxic to predators. Jasmonates also elicit the synthesis of volatile compounds that attract parasitoids, which are insects that spend their developing stages in or on another insect, and eventually kill their host. The plant may activate abscission of injured tissue if it is damaged beyond repair.
Summary
Plants respond to light by changes in morphology and activity. Irradiation by red light converts the photoreceptor phytochrome to its far-red light-absorbing form—Pfr. This form controls germination and flowering in response to length of day, as well as triggers photosynthesis in dormant plants or those that just emerged from the soil. Blue-light receptors, cryptochromes, and phototropins are responsible for phototropism. Amyloplasts, which contain heavy starch granules, sense gravity. Shoots exhibit negative gravitropism, whereas roots exhibit positive gravitropism. Plant hormones—naturally occurring compounds synthesized in small amounts—can act both in the cells that produce them and in distant tissues and organs. Auxins are responsible for apical dominance, root growth, directional growth toward light, and many other growth responses. Cytokinins stimulate cell division and counter apical dominance in shoots. Gibberellins inhibit dormancy of seeds and promote stem growth. Abscisic acid induces dormancy in seeds and buds, and protects plants from excessive water loss by promoting stomatal closure. Ethylene gas speeds up fruit ripening and dropping of leaves. Plants respond to touch by rapid movements (thigmotropy and thigmonasty) and slow differential growth (thigmomorphogenesis). Plants have evolved defense mechanisms against predators and pathogens. Physical barriers like bark and spines protect tender tissues. Plants also have chemical defenses, including toxic secondary metabolites and hormones, which elicit additional defense mechanisms.
Glossary
abscisic acid (ABA)
plant hormone that induces dormancy in seeds and other organs
abscission
physiological process that leads to the fall of a plant organ (such as leaf or petal drop)
auxin
plant hormone that influences cell elongation (in phototropism), gravitropism, apical dominance and root growth
chromophore
molecule that absorbs light
cryptochrome
protein that absorbs light in the blue and ultraviolet regions of the light spectrum
cytokinin
plant hormone that promotes cell division
ethylene
volatile plant hormone that is associated with fruit ripening, flower wilting, and leaf fall
gibberellin (GA)
plant hormone that stimulates shoot elongation, seed germination, and the maturation and dropping of fruit and flowers
jasmonates
small family of compounds derived from the fatty acid linoleic acid
negative gravitropism
growth away from Earth’s gravity
oligosaccharin
hormone important in plant defenses against bacterial and fungal infections
photomorphogenesis
growth and development of plants in response to light
photoperiodism
occurrence of plant processes, such as germination and flowering, according to the time of year
phototropin
blue-light receptor that promotes phototropism, stomatal opening and closing, and other responses that promote photosynthesis
phototropism
directional bending of a plant toward a light source
phytochrome
plant pigment protein that exists in two reversible forms (Pr and Pfr) and mediates morphologic changes in response to red light
positive gravitropism
growth toward Earth’s gravitational center
statolith
(also, amyloplast) plant organelle that contains heavy starch granules
strigolactone
hormone that promotes seed germination in some species and inhibits lateral apical development in the absence of auxins
thigmomorphogenesis
developmental response to touch
thigmonastic
directional growth of a plant independent of the direction in which contact is applied
thigmotropism
directional growth of a plant in response to constant contact | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.6%3A_Plant_Sensory_Systems_and_Responses.txt |
30.1: The Plant Body
Like animals, plants contain cells with organelles in which specific metabolic activities take place. Unlike animals, however, plants use energy from sunlight to form sugars during photosynthesis. In addition, plant cells have cell walls, plastids, and a large central vacuole: structures that are not found in animal cells. Each of these cellular structures plays a specific role in plant structure and function.
Review Questions
Plant regions of continuous growth are made up of ________.
1. dermal tissue
2. vascular tissue
3. meristematic tissue
4. permanent tissue
Answer
C
Which of the following is the major site of photosynthesis?
1. apical meristem
2. ground tissue
3. xylem cells
4. phloem cells
Answer
B
Free Response
What type of meristem is found only in monocots, such as lawn grasses? Explain how this type of meristematic tissue is beneficial in lawn grasses that are mowed each week.
Answer
Lawn grasses and other monocots have an intercalary meristem, which is a region of meristematic tissue at the base of the leaf blade. This is beneficial to the plant because it can continue to grow even when the tip of the plant is removed by grazing or mowing.
Which plant part is responsible for transporting water, minerals, and sugars to different parts of the plant? Name the two types of tissue that make up this overall tissue, and explain the role of each.
Answer
Vascular tissue transports water, minerals, and sugars throughout the plant. Vascular tissue is made up of xylem tissue and phloem tissue. Xylem tissue transports water and nutrients from the roots upward. Phloem tissue carries sugars from the sites of photosynthesis to the rest of the plant.
30.2: Stems
Plant stems, whether above or below ground, are characterized by the presence of nodes and internodes. Nodes are points of attachment for leaves, aerial roots, and flowers. The stem region between two nodes is called an internode. The stalk that extends from the stem to the base of the leaf is the petiole. An axillary bud is usually found in the axil—the area between the base of a leaf and the stem—where it can give rise to a branch or a flower.
Review Questions
Stem regions at which leaves are attached are called ________.
1. trichomes
2. lenticels
3. nodes
4. internodes
Answer
C
Which of the following cell types forms most of the inside of a plant?
1. meristem cells
2. collenchyma cells
3. sclerenchyma cells
4. parenchyma cells
Answer
D
Tracheids, vessel elements, sieve-tube cells, and companion cells are components of ________.
1. vascular tissue
2. meristematic tissue
3. ground tissue
4. dermal tissue
Answer
A
The primary growth of a plant is due to the action of the ________.
1. lateral meristem
2. vascular cambium
3. apical meristem
4. cork cambium
Answer
C
Which of the following is an example of secondary growth?
1. increase in length
2. increase in thickness or girth
3. increase in root hairs
4. increase in leaf number
Answer
B
Secondary growth in stems is usually seen in ________.
1. monocots
2. dicots
3. both monocots and dicots
4. neither monocots nor dicots
Answer
B
Free Response
Describe the roles played by stomata and guard cells. What would happen to a plant if these cells did not function correctly?
Answer
Stomata allow gases to enter and exit the plant. Guard cells regulate the opening and closing of stomata. If these cells did not function correctly, a plant could not get the carbon dioxide needed for photosynthesis, nor could it release the oxygen produced by photosynthesis.
Compare the structure and function of xylem to that of phloem.
Answer
Xylem is made up tracheids and vessel elements, which are cells that transport water and dissolved minerals and that are dead at maturity. Phloem is made up of sieve-tube cells and companion cells, which transport carbohydrates and are alive at maturity.
Explain the role of the cork cambium in woody plants.
Answer
In woody plants, the cork cambium is the outermost lateral meristem; it produces new cells towards the interior, which enables the plant to increase in girth. The cork cambium also produces cork cells towards the exterior, which protect the plant from physical damage while reducing water loss.
What is the function of lenticels?
Answer
In woody stems, lenticels allow internal cells to exchange gases with the outside atmosphere.
Besides the age of a tree, what additional information can annual rings reveal?
Answer
Annual rings can also indicate the climate conditions that prevailed during each growing season.
Give two examples of modified stems and explain how each example benefits the plant.
Answer
Answers will vary. Rhizomes, stolons, and runners can give rise to new plants. Corms, tubers, and bulbs can also produce new plants and can store food. Tendrils help a plant to climb, while thorns discourage herbivores.
30.3: Roots
The roots of seed plants have three major functions: anchoring the plant to the soil, absorbing water and minerals and transporting them upwards, and storing the products of photosynthesis. Some roots are modified to absorb moisture and exchange gases. Most roots are underground. Some plants, however, also have adventitious roots, which emerge above the ground from the shoot.
Review Questions
Roots that enable a plant to grow on another plant are called ________.
1. epiphytic roots
2. prop roots
3. adventitious roots
4. aerial roots
Answer
A
The ________ forces selective uptake of minerals in the root.
1. pericycle
2. epidermis
3. endodermis
4. root cap
Answer
C
Newly-formed root cells begin to form different cell types in the ________.
1. zone of elongation
2. zone of maturation
3. root meristem
4. zone of cell division
Answer
B
Free Response
Compare a tap root system with a fibrous root system. For each type, name a plant that provides a food in the human diet. Which type of root system is found in monocots? Which type of root system is found in dicots?
Answer
A tap root system has a single main root that grows down. A fibrous root system forms a dense network of roots that is closer to the soil surface. An example of a tap root system is a carrot. Grasses such as wheat, rice, and corn are examples of fibrous root systems. Fibrous root systems are found in monocots; tap root systems are found in dicots.
What might happen to a root if the pericycle disappeared?
Answer
The root would not be able to produce lateral roots.
30.4: Leaves
Leaves are the main sites for photosynthesis: the process by which plants synthesize food. Most leaves are usually green, due to the presence of chlorophyll in the leaf cells. However, some leaves may have different colors, caused by other plant pigments that mask the green chlorophyll. The thickness, shape, and size of leaves are adapted to the environment. Each variation helps a plant species maximize its chances of survival in a particular habitat.
Review Questions
The stalk of a leaf is known as the ________.
1. petiole
2. lamina
3. stipule
4. rachis
Answer
A
Leaflets are a characteristic of ________ leaves.
1. alternate
2. whorled
3. compound
4. opposite
Answer
C
Cells of the ________ contain chloroplasts.
1. epidermis
2. vascular tissue
3. stomata
4. mesophyll
Answer
D
Which of the following is most likely to be found in a desert environment?
1. broad leaves to capture sunlight
2. spines instead of leaves
3. needle-like leaves
4. wide, flat leaves that can float
Answer
B
Free Response
How do dicots differ from monocots in terms of leaf structure?
Answer
Monocots have leaves with parallel venation, and dicots have leaves with reticulate, net-like venation.
Describe an example of a plant with leaves that are adapted to cold temperatures.
Answer
Conifers such as spruce, fir, and pine have needle-shaped leaves with sunken stomata, helping to reduce water loss.
30.5: Transport of Water and Solutes in Plants
The structure of plant roots, stems, and leaves facilitates the transport of water, nutrients, and photosynthates throughout the plant. The phloem and xylem are the main tissues responsible for this movement. Water potential, evapotranspiration, and stomatal regulation influence how water and nutrients are transported in plants. To understand how these processes work, we must first understand the energetics of water potential.
Review Questions
When stomata open, what occurs?
1. Water vapor is lost to the external environment, increasing the rate of transpiration.
2. Water vapor is lost to the external environment, decreasing the rate of transpiration.
3. Water vapor enters the spaces in the mesophyll, increasing the rate of transpiration.
4. Water vapor enters the spaces in the mesophyll, increasing the rate of transpiration.
Answer
A
Which cells are responsible for the movement of photosynthates through a plant?
1. tracheids, vessel elements
2. tracheids, companion cells
3. vessel elements, companion cells
4. sieve-tube elements, companion cells
Answer
D
Free Response
The process of bulk flow transports fluids in a plant. Describe the two main bulk flow processes.
Answer
The process of bulk flow moves water up the xylem and moves photosynthates (solutes) up and down the phloem.
30.6: Plant Sensory Systems and Responses
Animals can respond to environmental factors by moving to a new location. Plants, however, are rooted in place and must respond to the surrounding environmental factors. Plants have sophisticated systems to detect and respond to light, gravity, temperature, and physical touch. Receptors sense environmental factors and relay the information to effector systems—often through intermediate chemical messengers—to bring about plant responses.
Review Questions
The main photoreceptor that triggers phototropism is a ________.
1. phytochrome
2. cryptochrome
3. phototropin
4. carotenoid
Answer
C
Phytochrome is a plant pigment protein that:
1. mediates plant infection
2. promotes plant growth
3. mediates morphological changes in response to red and far-red light
4. inhibits plant growth
Answer
C
A mutant plant has roots that grow in all directions. Which of the following organelles would you expect to be missing in the cell?
1. mitochondria
2. amyloplast
3. chloroplast
4. nucleus
Answer
B
After buying green bananas or unripe avocadoes, they can be kept in a brown bag to ripen. The hormone released by the fruit and trapped in the bag is probably:
1. abscisic acid
2. cytokinin
3. ethylene
4. gibberellic acid
Answer
C
A decrease in the level of which hormone releases seeds from dormancy?
1. abscisic acid
2. cytokinin
3. ethylene
4. gibberellic acid
Answer
A
A seedling germinating under a stone grows at an angle away from the stone and upward. This response to touch is called ________.
1. gravitropism
2. thigmonasty
3. thigmotropism
4. skototropism
Answer
C
Free Response
Owners and managers of plant nurseries have to plan lighting schedules for a long-day plant that will flower in February. What lighting periods will be most effective? What color of light should be chosen?
Answer
A long-day plant needs a higher proportion of the Pfr form to Pr form of phytochrome. The plant requires long periods of illumination with light enriched in the red range of the spectrum.
What are the major benefits of gravitropism for a germinating seedling?
Answer
Gravitropism will allow roots to dig deep into the soil to find water and minerals, whereas the seedling will grow towards light to enable photosynthesis.
Fruit and vegetable storage facilities are usually refrigerated and well ventilated. Why are these conditions advantageous?
Answer
Refrigeration slows chemical reactions, including fruit maturation. Ventilation removes the ethylene gas that speeds up fruit ripening.
Stomata close in response to bacterial infection. Why is this response a mechanism of defense for the plant? Which hormone is most likely to mediate this response?
Answer
To prevent further entry of pathogens, stomata close, even if they restrict entry of CO2. Some pathogens secrete virulence factors that inhibit the closing of stomata. Abscisic acid is the stress hormone responsible for inducing closing of stomata. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/30%3A_Plant_Form_and_Physiology/30.E%3A_Plant_Form_and_Physiology_%28Exercises%29.txt |
In order to grow and develop into mature, fruit-bearing plants, many requirements must be met and events must be coordinated. Seeds must germinate under the right conditions in the soil; therefore, temperature, moisture, and soil quality are important factors that play a role in germination and seedling development. Soil quality and climate are significant to plant distribution and growth.
• 31.0: Prelude to Soil and Plant Nutrition
This chapter will explore the complex dynamics between plants and soils, and the adaptations that plants have evolved to make better use of nutritional resources.
• 31.1: Nutritional Requirements of Plants
Plants are unique organisms that can absorb nutrients and water through their root system, as well as carbon dioxide from the atmosphere. Soil quality and climate are the major determinants of plant distribution and growth. The combination of soil nutrients, water, and carbon dioxide, along with sunlight, allows plants to grow.
• 31.2: The Soil
Soil is the outer loose layer that covers the surface of Earth. Soil quality is a major determinant, along with climate, of plant distribution and growth. Soil quality depends not only on the chemical composition of the soil, but also the topography (regional surface features) and the presence of living organisms. In agriculture, the history of the soil, such as the cultivating practices and previous crops, modify the characteristics and fertility of that soil.
• 31.3: Nutritional Adaptations of Plants
Plants obtain food in two different ways. Autotrophic plants can make their own food from inorganic raw materials, such as carbon dioxide and water, through photosynthesis in the presence of sunlight. Green plants are included in this group. Some plants, however, are heterotrophic: they are totally parasitic and lacking in chlorophyll. These plants, referred to as holo-parasitic plants, are unable to synthesize organic carbon and draw all of their nutrients from the host plant.
• 31.E: Soil and Plant Nutrition (Exercises)
31: Soil and Plant Nutrition
Cucurbitaceae is a family of plants first cultivated in Mesoamerica, although several species are native to North America. The family includes many edible species, such as squash and pumpkin, as well as inedible gourds. In order to grow and develop into mature, fruit-bearing plants, many requirements must be met and events must be coordinated. Seeds must germinate under the right conditions in the soil; therefore, temperature, moisture, and soil quality are important factors that play a role in germination and seedling development. Soil quality and climate are significant to plant distribution and growth. The young seedling will eventually grow into a mature plant, and the roots will absorb nutrients and water from the soil. At the same time, the aboveground parts of the plant will absorb carbon dioxide from the atmosphere and use energy from sunlight to produce organic compounds through photosynthesis. This chapter will explore the complex dynamics between plants and soils, and the adaptations that plants have evolved to make better use of nutritional resources.
31.1: Nutritional Requirements of Plants
Skills to Develop
• Describe how plants obtain nutrients
• List the elements and compounds required for proper plant nutrition
• Describe an essential nutrient
Plants are unique organisms that can absorb nutrients and water through their root system, as well as carbon dioxide from the atmosphere. Soil quality and climate are the major determinants of plant distribution and growth. The combination of soil nutrients, water, and carbon dioxide, along with sunlight, allows plants to grow.
The Chemical Composition of Plants
Since plants require nutrients in the form of elements such as carbon and potassium, it is important to understand the chemical composition of plants. The majority of volume in a plant cell is water; it typically comprises 80 to 90 percent of the plant’s total weight. Soil is the water source for land plants, and can be an abundant source of water, even if it appears dry. Plant roots absorb water from the soil through root hairs and transport it up to the leaves through the xylem. As water vapor is lost from the leaves, the process of transpiration and the polarity of water molecules (which enables them to form hydrogen bonds) draws more water from the roots up through the plant to the leaves (Figure \(1\)). Plants need water to support cell structure, for metabolic functions, to carry nutrients, and for photosynthesis.
Plant cells need essential substances, collectively called nutrients, to sustain life. Plant nutrients may be composed of either organic or inorganic compounds. An organic compound is a chemical compound that contains carbon, such as carbon dioxide obtained from the atmosphere. Carbon that was obtained from atmospheric CO2 composes the majority of the dry mass within most plants. An inorganic compound does not contain carbon and is not part of, or produced by, a living organism. Inorganic substances, which form the majority of the soil solution, are commonly called minerals: those required by plants include nitrogen (N) and potassium (K) for structure and regulation.
Essential Nutrients
Plants require only light, water and about 20 elements to support all their biochemical needs: these 20 elements are called essential nutrients. For an element to be regarded as essential, three criteria are required: 1) a plant cannot complete its life cycle without the element; 2) no other element can perform the function of the element; and 3) the element is directly involved in plant nutrition.
Table \(1\): Essential elements for plant growth
Macronutrients Micronutrients
Carbon (C) Iron (Fe)
Hydrogen (H) Manganese (Mn)
Oxygen (O) Boron (B)
Nitrogen (N) Molybdenum (Mo)
Phosphorus (P) Copper (Cu)
Potassium (K) Zinc (Zn)
Calcium (Ca) Chlorine (Cl)
Magnesium (Mg) Nickel (Ni)
Sulfur (S) Cobalt (Co)
Sodium (S)
Silicon (Si)
Macronutrients and Micronutrients
The essential elements can be divided into two groups: macronutrients and micronutrients. Nutrients that plants require in larger amounts are called macronutrients. About half of the essential elements are considered macronutrients: carbon, hydrogen, oxygen, nitrogen, phosphorus, potassium, calcium, magnesium and sulfur. The first of these macronutrients, carbon (C), is required to form carbohydrates, proteins, nucleic acids, and many other compounds; it is therefore present in all macromolecules. On average, the dry weight (excluding water) of a cell is 50 percent carbon. As shown in Figure \(2\), carbon is a key part of plant biomolecules.
The next most abundant element in plant cells is nitrogen (N); it is part of proteins and nucleic acids. Nitrogen is also used in the synthesis of some vitamins. Hydrogen and oxygen are macronutrients that are part of many organic compounds, and also form water. Oxygen is necessary for cellular respiration; plants use oxygen to store energy in the form of ATP. Phosphorus (P), another macromolecule, is necessary to synthesize nucleic acids and phospholipids. As part of ATP, phosphorus enables food energy to be converted into chemical energy through oxidative phosphorylation. Likewise, light energy is converted into chemical energy during photophosphorylation in photosynthesis, and into chemical energy to be extracted during respiration. Sulfur is part of certain amino acids, such as cysteine and methionine, and is present in several coenzymes. Sulfur also plays a role in photosynthesis as part of the electron transport chain, where hydrogen gradients play a key role in the conversion of light energy into ATP. Potassium (K) is important because of its role in regulating stomatal opening and closing. As the openings for gas exchange, stomata help maintain a healthy water balance; a potassium ion pump supports this process.
Magnesium (Mg) and calcium (Ca) are also important macronutrients. The role of calcium is twofold: to regulate nutrient transport, and to support many enzyme functions. Magnesium is important to the photosynthetic process. These minerals, along with the micronutrients, which are described below, also contribute to the plant’s ionic balance.
In addition to macronutrients, organisms require various elements in small amounts. These micronutrients, or trace elements, are present in very small quantities. They include boron (B), chlorine (Cl), manganese (Mn), iron (Fe), zinc (Zn), copper (Cu), molybdenum (Mo), nickel (Ni), silicon (Si), and sodium (Na).
Deficiencies in any of these nutrients—particularly the macronutrients—can adversely affect plant growth (Figure \(3\)). Depending on the specific nutrient, a lack can cause stunted growth, slow growth, or chlorosis (yellowing of the leaves). Extreme deficiencies may result in leaves showing signs of cell death.
Everyday Connection: Hydroponics
Hydroponics is a method of growing plants in a water-nutrient solution instead of soil. Since its advent, hydroponics has developed into a growing process that researchers often use. Scientists who are interested in studying plant nutrient deficiencies can use hydroponics to study the effects of different nutrient combinations under strictly controlled conditions. Hydroponics has also developed as a way to grow flowers, vegetables, and other crops in greenhouse environments. You might find hydroponically grown produce at your local grocery store. Today, many lettuces and tomatoes in your market have been hydroponically grown.
Summary
Plants can absorb inorganic nutrients and water through their root system, and carbon dioxide from the environment. The combination of organic compounds, along with water, carbon dioxide, and sunlight, produce the energy that allows plants to grow. Inorganic compounds form the majority of the soil solution. Plants access water though the soil. Water is absorbed by the plant root, transports nutrients throughout the plant, and maintains the structure of the plant. Essential elements are indispensable elements for plant growth. They are divided into macronutrients and micronutrients. The macronutrients plants require are carbon, nitrogen, hydrogen, oxygen, phosphorus, potassium, calcium, magnesium, and sulfur. Important micronutrients include iron, manganese, boron, molybdenum, copper, zinc, chlorine, nickel, cobalt, silicon and sodium.
Glossary
inorganic compound
chemical compound that does not contain carbon; it is not part of or produced by a living organism
macronutrient
nutrient that is required in large amounts for plant growth; carbon, hydrogen, oxygen, nitrogen, phosphorus, potassium, calcium, magnesium, and sulfur
micronutrient
nutrient required in small amounts; also called trace element
organic compound
chemical compound that contains carbon | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/31%3A_Soil_and_Plant_Nutrition/31.0%3A_Prelude_to_Soil_and_Plant_Nutrition.txt |
Skills to Develop
• Describe how soils are formed
• Explain soil composition
• Describe a soil profile
Plants obtain inorganic elements from the soil, which serves as a natural medium for land plants. Soil is the outer loose layer that covers the surface of Earth. Soil quality is a major determinant, along with climate, of plant distribution and growth. Soil quality depends not only on the chemical composition of the soil, but also the topography (regional surface features) and the presence of living organisms. In agriculture, the history of the soil, such as the cultivating practices and previous crops, modify the characteristics and fertility of that soil.
Soil develops very slowly over long periods of time, and its formation results from natural and environmental forces acting on mineral, rock, and organic compounds. Soils can be divided into two groups: organic soils are those that are formed from sedimentation and primarily composed of organic matter, while those that are formed from the weathering of rocks and are primarily composed of inorganic material are called mineral soils. Mineral soils are predominant in terrestrial ecosystems, where soils may be covered by water for part of the year or exposed to the atmosphere.
Soil Composition
Soil consists of these major components (Figure \(1\)):
• inorganic mineral matter, about 40 to 45 percent of the soil volume
• organic matter, about 5 percent of the soil volume
• water and air, about 50 percent of the soil volume
The amount of each of the four major components of soil depends on the amount of vegetation, soil compaction, and water present in the soil. A good healthy soil has sufficient air, water, minerals, and organic material to promote and sustain plant life.
Exercise \(1\)
Soil compaction can result when soil is compressed by heavy machinery or even foot traffic. How might this compaction change the soil composition?
Answer
The air content of the soil decreases.
The organic material of soil, called humus, is made up of microorganisms (dead and alive), and dead animals and plants in varying stages of decay. Humus improves soil structure and provides plants with water and minerals. The inorganic material of soil consists of rock, slowly broken down into smaller particles that vary in size. Soil particles that are 0.1 to 2 mm in diameter are sand. Soil particles between 0.002 and 0.1 mm are called silt, and even smaller particles, less than 0.002 mm in diameter, are called clay. Some soils have no dominant particle size and contain a mixture of sand, silt, and humus; these soils are called loams.
Soil Formation
Soil formation is the consequence of a combination of biological, physical, and chemical processes. Soil should ideally contain 50 percent solid material and 50 percent pore space. About one-half of the pore space should contain water, and the other half should contain air. The organic component of soil serves as a cementing agent, returns nutrients to the plant, allows soil to store moisture, makes soil tillable for farming, and provides energy for soil microorganisms. Most soil microorganisms—bacteria, algae, or fungi—are dormant in dry soil, but become active once moisture is available.
Soil distribution is not homogenous because its formation results in the production of layers; together, the vertical section of a soil is called the soil profile. Within the soil profile, soil scientists define zones called horizons. A horizon is a soil layer with distinct physical and chemical properties that differ from those of other layers. Five factors account for soil formation: parent material, climate, topography, biological factors, and time.
Parent Material
The organic and inorganic material in which soils form is the parent material. Mineral soils form directly from the weathering of bedrock, the solid rock that lies beneath the soil, and therefore, they have a similar composition to the original rock. Other soils form in materials that came from elsewhere, such as sand and glacial drift. Materials located in the depth of the soil are relatively unchanged compared with the deposited material. Sediments in rivers may have different characteristics, depending on whether the stream moves quickly or slowly. A fast-moving river could have sediments of rocks and sand, whereas a slow-moving river could have fine-textured material, such as clay.
Climate
Temperature, moisture, and wind cause different patterns of weathering and therefore affect soil characteristics. The presence of moisture and nutrients from weathering will also promote biological activity: a key component of a quality soil.
Topography
Regional surface features (familiarly called “the lay of the land”) can have a major influence on the characteristics and fertility of a soil. Topography affects water runoff, which strips away parent material and affects plant growth. Steeps soils are more prone to erosion and may be thinner than soils that are relatively flat or level.
Biological factors
The presence of living organisms greatly affects soil formation and structure. Animals and microorganisms can produce pores and crevices, and plant roots can penetrate into crevices to produce more fragmentation. Plant secretions promote the development of microorganisms around the root, in an area known as the rhizosphere. Additionally, leaves and other material that fall from plants decompose and contribute to soil composition.
Time
Time is an important factor in soil formation because soils develop over long periods. Soil formation is a dynamic process. Materials are deposited over time, decompose, and transform into other materials that can be used by living organisms or deposited onto the surface of the soil.
Physical Properties of the Soil
Soils are named and classified based on their horizons. The soil profile has four distinct layers: 1) O horizon; 2) A horizon; 3) B horizon, or subsoil; and 4) C horizon, or soil base (Figure \(2\)). The O horizon has freshly decomposing organic matter—humus—at its surface, with decomposed vegetation at its base. Humus enriches the soil with nutrients and enhances soil moisture retention. Topsoil—the top layer of soil—is usually two to three inches deep, but this depth can vary considerably. For instance, river deltas like the Mississippi River delta have deep layers of topsoil. Topsoil is rich in organic material; microbial processes occur there, and it is the “workhorse” of plant production. The A horizon consists of a mixture of organic material with inorganic products of weathering, and it is therefore the beginning of true mineral soil. This horizon is typically darkly colored because of the presence of organic matter. In this area, rainwater percolates through the soil and carries materials from the surface. The B horizon is an accumulation of mostly fine material that has moved downward, resulting in a dense layer in the soil. In some soils, the B horizon contains nodules or a layer of calcium carbonate. The C horizon, or soil base, includes the parent material, plus the organic and inorganic material that is broken down to form soil. The parent material may be either created in its natural place, or transported from elsewhere to its present location. Beneath the C horizon lies bedrock.
Exercise \(2\)
Which horizon is considered the topsoil, and which is considered the subsoil?
Answer
The A horizon is the topsoil, and the B horizon is subsoil.
Some soils may have additional layers, or lack one of these layers. The thickness of the layers is also variable, and depends on the factors that influence soil formation. In general, immature soils may have O, A, and C horizons, whereas mature soils may display all of these, plus additional layers (Figure \(3\)).
Career Connections: Soil Scientist
A soil scientist studies the biological components, physical and chemical properties, distribution, formation, and morphology of soils. Soil scientists need to have a strong background in physical and life sciences, plus a foundation in mathematics. They may work for federal or state agencies, academia, or the private sector. Their work may involve collecting data, carrying out research, interpreting results, inspecting soils, conducting soil surveys, and recommending soil management programs.
Many soil scientists work both in an office and in the field. According to the United States Department of Agriculture (USDA): “a soil scientist needs good observation skills to analyze and determine the characteristics of different types of soils. Soil types are complex and the geographical areas a soil scientist may survey are varied. Aerial photos or various satellite images are often used to research the areas. Computer skills and geographic information systems (GIS) help the scientist to analyze the multiple facets of geomorphology, topography, vegetation, and climate to discover the patterns left on the landscape.”1 Soil scientists play a key role in understanding the soil’s past, analyzing present conditions, and making recommendations for future soil-related practices.
Summary
Plants obtain mineral nutrients from the soil. Soil is the outer loose layer that covers the surface of Earth. Soil quality depends on the chemical composition of the soil, the topography, the presence of living organisms, the climate, and time. Agricultural practice and history may also modify the characteristics and fertility of soil. Soil consists of four major components: 1) inorganic mineral matter, 2) organic matter, 3) water and air, and 4) living matter. The organic material of soil is made of humus, which improves soil structure and provides water and minerals. Soil inorganic material consists of rock slowly broken down into smaller particles that vary in size, such as sand, silt, and loam.
Soil formation results from a combination of biological, physical, and chemical processes. Soil is not homogenous because its formation results in the production of layers called a soil profile. Factors that affect soil formation include: parent material, climate, topography, biological factors, and time. Soils are classified based on their horizons, soil particle size, and proportions. Most soils have four distinct horizons: O, A, B, and C.
Footnotes
1. 1 National Resources Conservation Service / United States Department of Agriculture. “Careers in Soil Science.” soils.usda.gov/education/facts/careers.html
Glossary
A horizon
consists of a mixture of organic material with inorganic products of weathering
B horizon
soil layer that is an accumulation of mostly fine material that has moved downward
bedrock
solid rock that lies beneath the soil
C horizon
layer of soil that contains the parent material, and the organic and inorganic material that is broken down to form soil; also known as the soil base
clay
soil particles that are less than 0.002 mm in diameter
horizon
soil layer with distinct physical and chemical properties, which differs from other layers depending on how and when it was formed
humus
organic material of soil; made up of microorganisms, dead animals and plants in varying stages of decay
loam
soil that has no dominant particle size
mineral soil
type of soil that is formed from the weathering of rocks and inorganic material; composed primarily of sand, silt, and clay
O horizon
layer of soil with humus at the surface and decomposed vegetation at the base
organic soil
type of soil that is formed from sedimentation; composed primarily of organic material
parent material
organic and inorganic material in which soils form
rhizosphere
area of soil affected by root secretions and microorganisms
sand
soil particles between 0.1–2 mm in diameter
silt
soil particles between 0.002 and 0.1 mm in diameter
soil profile
vertical section of a soil
soil
outer loose layer that covers the surface of Earth | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/31%3A_Soil_and_Plant_Nutrition/31.2%3A_The_Soil.txt |
Skills to Develop
• Understand the nutritional adaptations of plants
• Describe mycorrhizae
• Explain nitrogen fixation
Plants obtain food in two different ways. Autotrophic plants can make their own food from inorganic raw materials, such as carbon dioxide and water, through photosynthesis in the presence of sunlight. Green plants are included in this group. Some plants, however, are heterotrophic: they are totally parasitic and lacking in chlorophyll. These plants, referred to as holo-parasitic plants, are unable to synthesize organic carbon and draw all of their nutrients from the host plant.
Plants may also enlist the help of microbial partners in nutrient acquisition. Particular species of bacteria and fungi have evolved along with certain plants to create a mutualistic symbiotic relationship with roots. This improves the nutrition of both the plant and the microbe. The formation of nodules in legume plants and mycorrhization can be considered among the nutritional adaptations of plants. However, these are not the only type of adaptations that we may find; many plants have other adaptations that allow them to thrive under specific conditions.
Nitrogen Fixation: Root and Bacteria Interactions
Nitrogen is an important macronutrient because it is part of nucleic acids and proteins. Atmospheric nitrogen, which is the diatomic molecule $\ce{N2}$, or dinitrogen, is the largest pool of nitrogen in terrestrial ecosystems. However, plants cannot take advantage of this nitrogen because they do not have the necessary enzymes to convert it into biologically useful forms. However, nitrogen can be “fixed,” which means that it can be converted to ammonia ($\ce{NH3}$) through biological, physical, or chemical processes. As you have learned, biological nitrogen fixation (BNF) is the conversion of atmospheric nitrogen ($\ce{N2}$) into ammonia ($\ce{NH3}$), exclusively carried out by prokaryotes such as soil bacteria or cyanobacteria. Biological processes contribute 65 percent of the nitrogen used in agriculture. The following equation represents the process:
$\ce { N2 + 16 ATP + 8 e^{-} + 8 H^{+} \rightarrow 2 NH3 + 16 ADP + 16 P_i + H_2} \nonumber$
The most important source of BNF is the symbiotic interaction between soil bacteria and legume plants, including many crops important to humans (Figure $1$). The NH3 resulting from fixation can be transported into plant tissue and incorporated into amino acids, which are then made into plant proteins. Some legume seeds, such as soybeans and peanuts, contain high levels of protein, and serve among the most important agricultural sources of protein in the world.
Exercise $1$
Farmers often rotate corn (a cereal crop) and soy beans (a legume) planting a field with each crop in alternate seasons. What advantage might this crop rotation confer?
Answer
Soybeans are able to fix nitrogen in their roots, which are not harvested at the end of the growing season. The belowground nitrogen can be used in the next season by the corn.
Farmers often rotate corn (a cereal crop) and soy beans (a legume), planting a field with each crop in alternate seasons. What advantage might this crop rotation confer?
Soil bacteria, collectively called rhizobia, symbiotically interact with legume roots to form specialized structures called nodules, in which nitrogen fixation takes place. This process entails the reduction of atmospheric nitrogen to ammonia, by means of the enzyme nitrogenase. Therefore, using rhizobia is a natural and environmentally friendly way to fertilize plants, as opposed to chemical fertilization that uses a nonrenewable resource, such as natural gas. Through symbiotic nitrogen fixation, the plant benefits from using an endless source of nitrogen from the atmosphere. The process simultaneously contributes to soil fertility because the plant root system leaves behind some of the biologically available nitrogen. As in any symbiosis, both organisms benefit from the interaction: the plant obtains ammonia, and bacteria obtain carbon compounds generated through photosynthesis, as well as a protected niche in which to grow (Figure $2$).
Mycorrhizae: The Symbiotic Relationship between Fungi and Roots
A nutrient depletion zone can develop when there is rapid soil solution uptake, low nutrient concentration, low diffusion rate, or low soil moisture. These conditions are very common; therefore, most plants rely on fungi to facilitate the uptake of minerals from the soil. Fungi form symbiotic associations called mycorrhizae with plant roots, in which the fungi actually are integrated into the physical structure of the root. The fungi colonize the living root tissue during active plant growth.
Through mycorrhization, the plant obtains mainly phosphate and other minerals, such as zinc and copper, from the soil. The fungus obtains nutrients, such as sugars, from the plant root (Figure $3$). Mycorrhizae help increase the surface area of the plant root system because hyphae, which are narrow, can spread beyond the nutrient depletion zone. Hyphae can grow into small soil pores that allow access to phosphorus that would otherwise be unavailable to the plant. The beneficial effect on the plant is best observed in poor soils. The benefit to fungi is that they can obtain up to 20 percent of the total carbon accessed by plants. Mycorrhizae functions as a physical barrier to pathogens. It also provides an induction of generalized host defense mechanisms, and sometimes involves production of antibiotic compounds by the fungi.
There are two types of mycorrhizae: ectomycorrhizae and endomycorrhizae. Ectomycorrhizae form an extensive dense sheath around the roots, called a mantle. Hyphae from the fungi extend from the mantle into the soil, which increases the surface area for water and mineral absorption. This type of mycorrhizae is found in forest trees, especially conifers, birches, and oaks. Endomycorrhizae, also called arbuscular mycorrhizae, do not form a dense sheath over the root. Instead, the fungal mycelium is embedded within the root tissue. Endomycorrhizae are found in the roots of more than 80 percent of terrestrial plants.
Nutrients from Other Sources
Some plants cannot produce their own food and must obtain their nutrition from outside sources. This may occur with plants that are parasitic or saprophytic. Some plants are mutualistic symbionts, epiphytes, or insectivorous.
Plant Parasites
A parasitic plant depends on its host for survival. Some parasitic plants have no leaves. An example of this is the dodder (Figure $4$), which has a weak, cylindrical stem that coils around the host and forms suckers. From these suckers, cells invade the host stem and grow to connect with the vascular bundles of the host. The parasitic plant obtains water and nutrients through these connections. The plant is a total parasite (a holoparasite) because it is completely dependent on its host. Other parasitic plants (hemiparasites) are fully photosynthetic and only use the host for water and minerals. There are about 4,100 species of parasitic plants.
Saprophytes
A saprophyte is a plant that does not have chlorophyll and gets its food from dead matter, similar to bacteria and fungi (note that fungi are often called saprophytes, which is incorrect, because fungi are not plants). Plants like these use enzymes to convert organic food materials into simpler forms from which they can absorb nutrients (Figure $5$). Most saprophytes do not directly digest dead matter: instead, they parasitize fungi that digest dead matter, or are mycorrhizal, ultimately obtaining photosynthate from a fungus that derived photosynthate from its host. Saprophytic plants are uncommon; only a few species are described.
Symbionts
A symbiont is a plant in a symbiotic relationship, with special adaptations such as mycorrhizae or nodule formation. Fungi also form symbiotic associations with cyanobacteria and green algae (called lichens). Lichens can sometimes be seen as colorful growths on the surface of rocks and trees (Figure $6$). The algal partner (phycobiont) makes food autotrophically, some of which it shares with the fungus; the fungal partner (mycobiont) absorbs water and minerals from the environment, which are made available to the green alga. If one partner was separated from the other, they would both die.
Epiphytes
An epiphyte is a plant that grows on other plants, but is not dependent upon the other plant for nutrition (Figure $7$). Epiphytes have two types of roots: clinging aerial roots, which absorb nutrients from humus that accumulates in the crevices of trees; and aerial roots, which absorb moisture from the atmosphere.
Insectivorous Plants
An insectivorous plant has specialized leaves to attract and digest insects. The Venus flytrap is popularly known for its insectivorous mode of nutrition, and has leaves that work as traps (Figure $8$). The minerals it obtains from prey compensate for those lacking in the boggy (low pH) soil of its native North Carolina coastal plains. There are three sensitive hairs in the center of each half of each leaf. The edges of each leaf are covered with long spines. Nectar secreted by the plant attracts flies to the leaf. When a fly touches the sensory hairs, the leaf immediately closes. Next, fluids and enzymes break down the prey and minerals are absorbed by the leaf. Since this plant is popular in the horticultural trade, it is threatened in its original habitat.
Summary
Atmospheric nitrogen is the largest pool of available nitrogen in terrestrial ecosystems. However, plants cannot use this nitrogen because they do not have the necessary enzymes. Biological nitrogen fixation (BNF) is the conversion of atmospheric nitrogen to ammonia. The most important source of BNF is the symbiotic interaction between soil bacteria and legumes. The bacteria form nodules on the legume’s roots in which nitrogen fixation takes place. Fungi form symbiotic associations (mycorrhizae) with plants, becoming integrated into the physical structure of the root. Through mycorrhization, the plant obtains minerals from the soil and the fungus obtains photosynthate from the plant root. Ectomycorrhizae form an extensive dense sheath around the root, while endomycorrhizae are embedded within the root tissue. Some plants—parasites, saprophytes, symbionts, epiphytes, and insectivores—have evolved adaptations to obtain their organic or mineral nutrition from various sources.
Glossary
epiphyte
plant that grows on other plants but is not dependent upon other plants for nutrition
insectivorous plant
plant that has specialized leaves to attract and digest insects
nitrogenase
enzyme that is responsible for the reduction of atmospheric nitrogen to ammonia
nodules
specialized structures that contain Rhizobia bacteria where nitrogen fixation takes place
parasitic plant
plant that is dependent on its host for survival
rhizobia
soil bacteria that symbiotically interact with legume roots to form nodules and fix nitrogen
saprophyte
plant that does not have chlorophyll and gets its food from dead matter
symbiont
plant in a symbiotic relationship with bacteria or fungi | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/31%3A_Soil_and_Plant_Nutrition/31.3%3A_Nutritional_Adaptations_of_Plants.txt |
31.1: Nutritional Requirements of Plants
Plants are unique organisms that can absorb nutrients and water through their root system, as well as carbon dioxide from the atmosphere. Soil quality and climate are the major determinants of plant distribution and growth. The combination of soil nutrients, water, and carbon dioxide, along with sunlight, allows plants to grow.
Review Questions
For an element to be regarded as essential, all of the following criteria must be met, except:
1. No other element can perform the function.
2. The element is directly involved in plant nutrition.
3. The element is inorganic.
4. The plant cannot complete its lifecycle without the element.
Answer
C
The nutrient that is part of carbohydrates, proteins, and nucleic acids, and that forms biomolecules, is ________.
1. nitrogen
2. carbon
3. magnesium
4. iron
Answer
B
Most ________ are necessary for enzyme function.
1. micronutrients
2. macronutrients
3. biomolecules
4. essential nutrients
Answer
A
What is the main water source for land plants?
1. rain
2. soil
3. biomolecules
4. essential nutrients
Answer
B
Free Response
What type of plant problems result from nitrogen and calcium deficiencies?
Answer
Deficiencies in these nutrients could result in stunted growth, slow growth, and chlorosis.
What did the van Helmont experiment show?
Answer
van Helmont showed that plants do not consume soil, which is correct. He also thought that plant growth and increased weight resulted from the intake of water, a conclusion that has since been disproven.
List two essential macronutrients and two essential nutrients.
Answer
Answers may vary. Essential macronutrients include carbon, hydrogen, oxygen, nitrogen, phosphorus, potassium, calcium, magnesium, and sulfur. Essential micronutrients include iron, manganese, boron, molybdenum, copper, zinc, chlorine, nickel, cobalt, sodium, and silicon.
31.2: The Soil
Soil is the outer loose layer that covers the surface of Earth. Soil quality is a major determinant, along with climate, of plant distribution and growth. Soil quality depends not only on the chemical composition of the soil, but also the topography (regional surface features) and the presence of living organisms. In agriculture, the history of the soil, such as the cultivating practices and previous crops, modify the characteristics and fertility of that soil.
Review Questions
Which factors affect soil quality?
1. chemical composition
2. history of the soil
3. presence of living organisms and topography
4. all of the above
Answer
D
Soil particles that are 0.1 to 2 mm in diameter are called ________.
1. sand
2. silt
3. clay
4. loam
Answer
A
A soil consists of layers called ________ that taken together are called a ________.
1. soil profiles : horizon
2. horizons : soil profile
3. horizons : humus
4. humus : soil profile
Answer
B
What is the term used to describe the solid rock that lies beneath the soil?
1. sand
2. bedrock
3. clay
4. loam
Answer
B
Free Response
Describe the main differences between a mineral soil and an organic soil.
Answer
A mineral soil forms from the weathering of rocks; it is inorganic material. An organic soil is formed from sedimentation; it mostly consists of humus.
Name and briefly explain the factors that affect soil formation.
Answer
Parent material, climate, topography, biological factors, and time affect soil formation. Parent material is the material in which soils form. Climate describes how temperature, moisture, and wind cause different patterns of weathering, influencing the characteristics of the soil. Topography affects the characteristics and fertility of a soil. Biological factors include the presence of living organisms that greatly affect soil formation. Processes such as freezing and thawing may produce cracks in rocks; plant roots can penetrate these crevices and produce more fragmentation. Time affects soil because soil develops over long periods.
Describe how topography influences the characteristics and fertility of a soil.
Answer
Topography affects water runoff, which strips away parent material and affects plant growth. Steeps soils are more prone to erosion and may be thinner than soils that are on level surfaces.
31.3: Nutritional Adaptations of Plants
Plants obtain food in two different ways. Autotrophic plants can make their own food from inorganic raw materials, such as carbon dioxide and water, through photosynthesis in the presence of sunlight. Green plants are included in this group. Some plants, however, are heterotrophic: they are totally parasitic and lacking in chlorophyll. These plants, referred to as holo-parasitic plants, are unable to synthesize organic carbon and draw all of their nutrients from the host plant.
Review Questions
Which process produces an inorganic compound that plants can easily use?
1. photosynthesis
2. nitrogen fixation
3. mycorrhization
4. Calvin cycle
Answer
B
Through mycorrhization, a plant obtains important nutrients such as ________.
1. phosphorus, zinc, and copper
2. phosphorus, zinc, and calcium
3. nickel, calcium, and zinc
4. all of the above
Answer
A
What term describes a plant that requires nutrition from a living host plant?
1. parasite
2. saprophyte
3. epiphyte
4. insectivorous
Answer
A
What is the term for the symbiotic association between fungi and cyanobacteria?
1. lichen
2. mycorrhizae
3. epiphyte
4. nitrogen-fixing nodule
Answer
A
Free Response
Why is biological nitrogen fixation an environmentally friendly way of fertilizing plants?
Answer
Because it is natural and does not require use of a nonrenewable resource, such as natural gas.
What is the main difference, from an energy point of view, between photosynthesis and biological nitrogen fixation?
Answer
Photosynthesis harvests and stores energy, whereas biological nitrogen fixation requires energy.
Why is a root nodule a nutritional adaptation of a plant?
Answer
A nodule results from the symbiosis between a plant and bacterium. Within nodules, the process of nitrogen fixation allows the plant to obtain nitrogen from the air. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/31%3A_Soil_and_Plant_Nutrition/31.E%3A_Soil_and_Plant_Nutrition_%28Exercises%29.txt |
Plant reproduction in plants can be accomplished via either sexual or asexual mechanisms. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from the parent or parents. Asexual reproduction produces new individuals without the fusion of gametes, genetically identical to the parent plants and each other, except when mutations occur. In seed plants, the offspring can be packaged in a protective seed, which is used as an agent of dispersal.
• 32.0: Prelude to Plant Reproduction
Plants have evolved different reproductive strategies for the continuation of their species. Some plants reproduce sexually, and others asexually, in contrast to animal species, which rely almost exclusively on sexual reproduction. Plant sexual reproduction usually depends on pollinating agents, while asexual reproduction is independent of these agents. Flowers are often the showiest or most strongly scented part of plants.
• 32.1: Reproductive Development and Structure
Sexual reproduction takes place with slight variations in different groups of plants. Plants have two distinct stages in their lifecycle: the gametophyte stage and the sporophyte stage. The haploid gametophyte produces the male and female gametes by mitosis in distinct multicellular structures. Fusion of the male and females gametes forms the diploid zygote, which develops into the sporophyte.
• 32.2: Pollination and Fertilization
Pollination takes two forms: self-pollination and cross-pollination. Self-pollination occurs when the pollen from the anther is deposited on the stigma of the same flower, or another flower on the same plant. Cross-pollination is the transfer of pollen from the anther of one flower to the stigma of another flower on a different individual of the same species. Self-pollination occurs in flowers where the stamen and carpel mature at the same time.
• 32.3: Asexual Reproduction
Many plants are able to propagate themselves using asexual reproduction. This method does not require the investment required to produce a flower, attract pollinators, or find a means of seed dispersal. Asexual reproduction produces plants that are genetically identical to the parent plant because no mixing of male and female gametes takes place. Traditionally, these plants survive well under stable environmental conditions when compared with plants produced from sexual reproduction.
• 32.E: Plant Reproduction (Exercises)
32: Plant Reproduction
Plants have evolved different reproductive strategies for the continuation of their species. Some plants reproduce sexually, and others asexually, in contrast to animal species, which rely almost exclusively on sexual reproduction. Plant sexual reproduction usually depends on pollinating agents, while asexual reproduction is independent of these agents. Flowers are often the showiest or most strongly scented part of plants. With their bright colors, fragrances, and interesting shapes and sizes, flowers attract insects, birds, and animals to serve their pollination needs. Other plants pollinate via wind or water; still others self-pollinate.
32.1: Reproductive Development and Structure
Skills to Develop
• Describe the two stages of a plant’s lifecycle
• Compare and contrast male and female gametophytes and explain how they form in angiosperms
• Describe the reproductive structures of a plant
• Describe the components of a complete flower
• Describe the development of microsporangium and megasporangium in gymnosperms
Sexual reproduction takes place with slight variations in different groups of plants. Plants have two distinct stages in their lifecycle: the gametophyte stage and the sporophyte stage. The haploid gametophyte produces the male and female gametes by mitosis in distinct multicellular structures. Fusion of the male and females gametes forms the diploid zygote, which develops into the sporophyte. After reaching maturity, the diploid sporophyte produces spores by meiosis, which in turn divide by mitosis to produce the haploid gametophyte. The new gametophyte produces gametes, and the cycle continues. This is the alternation of generations, and is typical of plant reproduction (Figure \(1\)).
The life cycle of higher plants is dominated by the sporophyte stage, with the gametophyte borne on the sporophyte. In ferns, the gametophyte is free-living and very distinct in structure from the diploid sporophyte. In bryophytes, such as mosses, the haploid gametophyte is more developed than the sporophyte.
During the vegetative phase of growth, plants increase in size and produce a shoot system and a root system. As they enter the reproductive phase, some of the branches start to bear flowers. Many flowers are borne singly, whereas some are borne in clusters. The flower is borne on a stalk known as a receptacle. Flower shape, color, and size are unique to each species, and are often used by taxonomists to classify plants.
Sexual Reproduction in Angiosperms
The lifecycle of angiosperms follows the alternation of generations explained previously. The haploid gametophyte alternates with the diploid sporophyte during the sexual reproduction process of angiosperms. Flowers contain the plant’s reproductive structures.
Flower Structure
A typical flower has four main parts—or whorls—known as the calyx, corolla, androecium, and gynoecium (Figure \(2\)). The outermost whorl of the flower has green, leafy structures known as sepals. The sepals, collectively called the calyx, help to protect the unopened bud. The second whorl is comprised of petals—usually, brightly colored—collectively called the corolla. The number of sepals and petals varies depending on whether the plant is a monocot or dicot. In monocots, petals usually number three or multiples of three; in dicots, the number of petals is four or five, or multiples of four and five. Together, the calyx and corolla are known as the perianth. The third whorl contains the male reproductive structures and is known as the androecium. The androecium has stamens with anthers that contain the microsporangia. The innermost group of structures in the flower is the gynoecium, or the female reproductive component(s). The carpel is the individual unit of the gynoecium and has a stigma, style, and ovary. A flower may have one or multiple carpels.
Art Connection
If the anther is missing, what type of reproductive structure will the flower be unable to produce? What term is used to describe an incomplete flower lacking the androecium? What term describes an incomplete flower lacking a gynoecium?
If all four whorls (the calyx, corolla, androecium, and gynoecium) are present, the flower is described as complete. If any of the four parts is missing, the flower is known as incomplete. Flowers that contain both an androecium and a gynoecium are called perfect, androgynous or hermaphrodites. There are two types of incomplete flowers: staminate flowers contain only an androecium, and carpellate flowers have only a gynoecium (Figure \(3\)).
If both male and female flowers are borne on the same plant, the species is called monoecious (meaning “one home”): examples are corn and pea. Species with male and female flowers borne on separate plants are termed dioecious, or “two homes,” examples of which are C. papaya and Cannabis. The ovary, which may contain one or multiple ovules, may be placed above other flower parts, which is referred to as superior; or, it may be placed below the other flower parts, referred to as inferior (Figure \(4\)).
Male Gametophyte (The Pollen Grain)
The male gametophyte develops and reaches maturity in an immature anther. In a plant’s male reproductive organs, development of pollen takes place in a structure known as the microsporangium (Figure \(5\)). The microsporangia, which are usually bi-lobed, are pollen sacs in which the microspores develop into pollen grains. These are found in the anther, which is at the end of the stamen—the long filament that supports the anther.
Within the microsporangium, the microspore mother cell divides by meiosis to give rise to four microspores, each of which will ultimately form a pollen grain (Figure \(6\)). An inner layer of cells, known as the tapetum, provides nutrition to the developing microspores and contributes key components to the pollen wall. Mature pollen grains contain two cells: a generative cell and a pollen tube cell. The generative cell is contained within the larger pollen tube cell. Upon germination, the tube cell forms the pollen tube through which the generative cell migrates to enter the ovary. During its transit inside the pollen tube, the generative cell divides to form two male gametes (sperm cells). Upon maturity, the microsporangia burst, releasing the pollen grains from the anther.
Each pollen grain has two coverings: the exine (thicker, outer layer) and the intine (Figure \(6\)). The exine contains sporopollenin, a complex waterproofing substance supplied by the tapetal cells. Sporopollenin allows the pollen to survive under unfavorable conditions and to be carried by wind, water, or biological agents without undergoing damage.
Female Gametophyte (The Embryo Sac)
While the details may vary between species, the overall development of the female gametophyte has two distinct phases. First, in the process of megasporogenesis, a single cell in the diploid megasporangium—an area of tissue in the ovules—undergoes meiosis to produce four megaspores, only one of which survives. During the second phase, megagametogenesis, the surviving haploid megaspore undergoes mitosis to produce an eight-nucleate, seven-cell female gametophyte, also known as the megagametophyte or embryo sac. Two of the nuclei—the polar nuclei—move to the equator and fuse, forming a single, diploid central cell. This central cell later fuses with a sperm to form the triploid endosperm. Three nuclei position themselves on the end of the embryo sac opposite the micropyle and develop into the antipodal cells, which later degenerate. The nucleus closest to the micropyle becomes the female gamete, or egg cell, and the two adjacent nuclei develop into synergid cells (Figure \(7\)). The synergids help guide the pollen tube for successful fertilization, after which they disintegrate. Once fertilization is complete, the resulting diploid zygote develops into the embryo, and the fertilized ovule forms the other tissues of the seed.
A double-layered integument protects the megasporangium and, later, the embryo sac. The integument will develop into the seed coat after fertilization and protect the entire seed. The ovule wall will become part of the fruit. The integuments, while protecting the megasporangium, do not enclose it completely, but leave an opening called the micropyle. The micropyle allows the pollen tube to enter the female gametophyte for fertilization.
Art Connection
An embryo sac is missing the synergids. What specific impact would you expect this to have on fertilization?
1. The pollen tube will be unable to form.
2. The pollen tube will form but will not be guided toward the egg.
3. Fertilization will not occur because the synergid is the egg.
4. Fertilization will occur but the embryo will not be able to grow.
Sexual Reproduction in Gymnosperms
As with angiosperms, the lifecycle of a gymnosperm is also characterized by alternation of generations. In conifers such as pines, the green leafy part of the plant is the sporophyte, and the cones contain the male and female gametophytes (Figure \(8\)). The female cones are larger than the male cones and are positioned towards the top of the tree; the small, male cones are located in the lower region of the tree. Because the pollen is shed and blown by the wind, this arrangement makes it difficult for a gymnosperm to self-pollinate.
Male Gametophyte
A male cone has a central axis on which bracts, a type of modified leaf, are attached. The bracts are known as microsporophylls (Figure \(9\)) and are the sites where microspores will develop. The microspores develop inside the microsporangium. Within the microsporangium, cells known as microsporocytes divide by meiosis to produce four haploid microspores. Further mitosis of the microspore produces two nuclei: the generative nucleus, and the tube nucleus. Upon maturity, the male gametophyte (pollen) is released from the male cones and is carried by the wind to land on the female cone.
Link to Learning
Watch this video to see a cedar releasing its pollen in the wind.
Female Gametophyte
The female cone also has a central axis on which bracts known as megasporophylls (Figure \(9\)) are present. In the female cone, megaspore mother cells are present in the megasporangium. The megaspore mother cell divides by meiosis to produce four haploid megaspores. One of the megaspores divides to form the multicellular female gametophyte, while the others divide to form the rest of the structure. The female gametophyte is contained within a structure called the archegonium.
Reproductive Process
Upon landing on the female cone, the tube cell of the pollen forms the pollen tube, through which the generative cell migrates towards the female gametophyte through the micropyle. It takes approximately one year for the pollen tube to grow and migrate towards the female gametophyte. The male gametophyte containing the generative cell splits into two sperm nuclei, one of which fuses with the egg, while the other degenerates. After fertilization of the egg, the diploid zygote is formed, which divides by mitosis to form the embryo. The scales of the cones are closed during development of the seed. The seed is covered by a seed coat, which is derived from the female sporophyte. Seed development takes another one to two years. Once the seed is ready to be dispersed, the bracts of the female cones open to allow the dispersal of seed; no fruit formation takes place because gymnosperm seeds have no covering.
Angiosperms versus Gymnosperms
Gymnosperm reproduction differs from that of angiosperms in several ways (Figure \(10\)). In angiosperms, the female gametophyte exists in an enclosed structure—the ovule—which is within the ovary; in gymnosperms, the female gametophyte is present on exposed bracts of the female cone. Double fertilization is a key event in the lifecycle of angiosperms, but is completely absent in gymnosperms. The male and female gametophyte structures are present on separate male and female cones in gymnosperms, whereas in angiosperms, they are a part of the flower. Lastly, wind plays an important role in pollination in gymnosperms because pollen is blown by the wind to land on the female cones. Although many angiosperms are also wind-pollinated, animal pollination is more common.
Link to Learning
Visit this website to view an animation of the double fertilization process of angiosperms.
Summary
The flower contains the reproductive structures of a plant. All complete flowers contain four whorls: the calyx, corolla, androecium, and gynoecium. The stamens are made up of anthers, in which pollen grains are produced, and a supportive strand called the filament. The pollen contains two cells— a generative cell and a tube cell—and is covered by two layers called the intine and the exine. The carpels, which are the female reproductive structures, consist of the stigma, style, and ovary. The female gametophyte is formed from mitotic divisions of the megaspore, forming an eight-nuclei ovule sac. This is covered by a layer known as the integument. The integument contains an opening called the micropyle, through which the pollen tube enters the embryo sac.
The diploid sporophyte of angiosperms and gymnosperms is the conspicuous and long-lived stage of the life cycle. The sporophytes differentiate specialized reproductive structures called sporangia, which are dedicated to the production of spores. The microsporangium contains microspore mother cells, which divide by meiosis to produce haploid microspores. The microspores develop into male gametophytes that are released as pollen. The megasporangium contains megaspore mother cells, which divide by meiosis to produce haploid megaspores. A megaspore develops into a female gametophyte containing a haploid egg. A new diploid sporophyte is formed when a male gamete from a pollen grain enters the ovule sac and fertilizes this egg.
Art Connections
Figure \(2\): If the anther is missing, what type of reproductive structure will the flower be unable to produce? What term is used to describe a flower that is normally lacking the androecium? What term describes a flower lacking a gynoecium?
Answer
Pollen (or sperm); carpellate; staminate.
Figure \(7\): An embryo sac is missing the synergids. What specific impact would you expect this to have on fertilization?
1. The pollen tube will be unable to form.
2. The pollen tube will form but will not be guided toward the egg.
3. Fertilization will not occur because the synergid is the egg.
4. Fertilization will occur but the embryo will not be able to grow.
Answer
B: The pollen tube will form but will not be guided toward the egg.
Glossary
androecium
sum of all the stamens in a flower
antipodals
the three cells away from the micropyle
exine
outermost covering of pollen
gametophyte
multicellular stage of the plant that gives rise to haploid gametes or spores
gynoecium
the sum of all the carpels in a flower
intine
inner lining of the pollen
megagametogenesis
second phase of female gametophyte development, during which the surviving haploid megaspore undergoes mitosis to produce an eight-nucleate, seven-cell female gametophyte, also known as the megagametophyte or embryo sac.
megasporangium
tissue found in the ovary that gives rise to the female gamete or egg
megasporogenesis
first phase of female gametophyte development, during which a single cell in the diploid megasporangium undergoes meiosis to produce four megaspores, only one of which survives
megasporophyll
bract (a type of modified leaf) on the central axis of a female gametophyte
micropyle
opening on the ovule sac through which the pollen tube can gain entry
microsporangium
tissue that gives rise to the microspores or the pollen grain
microsporophyll
central axis of a male cone on which bracts (a type of modified leaf) are attached
perianth
(also, petal or sepal) part of the flower consisting of the calyx and/or corolla; forms the outer envelope of the flower
polar nuclei
found in the ovule sac; fusion with one sperm cell forms the endosperm
sporophyte
multicellular diploid stage in plants that is formed after the fusion of male and female gametes
synergid
type of cell found in the ovule sac that secretes chemicals to guide the pollen tube towards the egg | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/32%3A_Plant_Reproduction/32.0%3A_Prelude_to_Plant_Reproduction.txt |
Skills to Develop
• Describe what must occur for plant fertilization
• Explain cross-pollination and the ways in which it takes place
• Describe the process that leads to the development of a seed
• Define double fertilization
In angiosperms, pollination is defined as the placement or transfer of pollen from the anther to the stigma of the same flower or another flower. In gymnosperms, pollination involves pollen transfer from the male cone to the female cone. Upon transfer, the pollen germinates to form the pollen tube and the sperm for fertilizing the egg. Pollination has been well studied since the time of Gregor Mendel. Mendel successfully carried out self- as well as cross-pollination in garden peas while studying how characteristics were passed on from one generation to the next. Today’s crops are a result of plant breeding, which employs artificial selection to produce the present-day cultivars. A case in point is today's corn, which is a result of years of breeding that started with its ancestor, teosinte. The teosinte that the ancient Mayans originally began cultivating had tiny seeds—vastly different from today’s relatively giant ears of corn. Interestingly, though these two plants appear to be entirely different, the genetic difference between them is miniscule.
Pollination takes two forms: self-pollination and cross-pollination. Self-pollination occurs when the pollen from the anther is deposited on the stigma of the same flower, or another flower on the same plant. Cross-pollination is the transfer of pollen from the anther of one flower to the stigma of another flower on a different individual of the same species. Self-pollination occurs in flowers where the stamen and carpel mature at the same time, and are positioned so that the pollen can land on the flower’s stigma. This method of pollination does not require an investment from the plant to provide nectar and pollen as food for pollinators.
Link to Learning
Explore this interactive website to review self-pollination and cross-pollination: http://passel.unl.edu/pages/animatio...erbreeding.swf
Living species are designed to ensure survival of their progeny; those that fail become extinct. Genetic diversity is therefore required so that in changing environmental or stress conditions, some of the progeny can survive. Self-pollination leads to the production of plants with less genetic diversity, since genetic material from the same plant is used to form gametes, and eventually, the zygote. In contrast, cross-pollination—or out-crossing—leads to greater genetic diversity because the microgametophyte and megagametophyte are derived from different plants.
Because cross-pollination allows for more genetic diversity, plants have developed many ways to avoid self-pollination. In some species, the pollen and the ovary mature at different times. These flowers make self-pollination nearly impossible. By the time pollen matures and has been shed, the stigma of this flower is mature and can only be pollinated by pollen from another flower. Some flowers have developed physical features that prevent self-pollination. The primrose is one such flower. Primroses have evolved two flower types with differences in anther and stigma length: the pin-eyed flower has anthers positioned at the pollen tube’s halfway point, and the thrum-eyed flower’s stigma is likewise located at the halfway point. Insects easily cross-pollinate while seeking the nectar at the bottom of the pollen tube. This phenomenon is also known as heterostyly. Many plants, such as cucumber, have male and female flowers located on different parts of the plant, thus making self-pollination difficult. In yet other species, the male and female flowers are borne on different plants (dioecious). All of these are barriers to self-pollination; therefore, the plants depend on pollinators to transfer pollen. The majority of pollinators are biotic agents such as insects (like bees, flies, and butterflies), bats, birds, and other animals. Other plant species are pollinated by abiotic agents, such as wind and water.
Everyday Connection: Incompatibility Genes in Flowers
In recent decades, incompatibility genes—which prevent pollen from germinating or growing into the stigma of a flower—have been discovered in many angiosperm species. If plants do not have compatible genes, the pollen tube stops growing. Self-incompatibility is controlled by the S (sterility) locus. Pollen tubes have to grow through the tissue of the stigma and style before they can enter the ovule. The carpel is selective in the type of pollen it allows to grow inside. The interaction is primarily between the pollen and the stigma epidermal cells. In some plants, like cabbage, the pollen is rejected at the surface of the stigma, and the unwanted pollen does not germinate. In other plants, pollen tube germination is arrested after growing one-third the length of the style, leading to pollen tube death. Pollen tube death is due either to apoptosis (programmed cell death) or to degradation of pollen tube RNA. The degradation results from the activity of a ribonuclease encoded by the S locus. The ribonuclease is secreted from the cells of the style in the extracellular matrix, which lies alongside the growing pollen tube.
In summary, self-incompatibility is a mechanism that prevents self-fertilization in many flowering plant species. The working of this self-incompatibility mechanism has important consequences for plant breeders because it inhibits the production of inbred and hybrid plants.
Pollination by Insects
Bees are perhaps the most important pollinator of many garden plants and most commercial fruit trees (Figure \(1\)). The most common species of bees are bumblebees and honeybees. Since bees cannot see the color red, bee-pollinated flowers usually have shades of blue, yellow, or other colors. Bees collect energy-rich pollen or nectar for their survival and energy needs. They visit flowers that are open during the day, are brightly colored, have a strong aroma or scent, and have a tubular shape, typically with the presence of a nectar guide. A nectar guide includes regions on the flower petals that are visible only to bees, and not to humans; it helps to guide bees to the center of the flower, thus making the pollination process more efficient. The pollen sticks to the bees’ fuzzy hair, and when the bee visits another flower, some of the pollen is transferred to the second flower. Recently, there have been many reports about the declining population of honeybees. Many flowers will remain unpollinated and not bear seed if honeybees disappear. The impact on commercial fruit growers could be devastating.
Many flies are attracted to flowers that have a decaying smell or an odor of rotting flesh. These flowers, which produce nectar, usually have dull colors, such as brown or purple. They are found on the corpse flower or voodoo lily (Amorphophallus), dragon arum (Dracunculus), and carrion flower (Stapleia, Rafflesia). The nectar provides energy, whereas the pollen provides protein. Wasps are also important insect pollinators, and pollinate many species of figs.
Butterflies, such as the monarch, pollinate many garden flowers and wildflowers, which usually occur in clusters. These flowers are brightly colored, have a strong fragrance, are open during the day, and have nectar guides to make access to nectar easier. The pollen is picked up and carried on the butterfly’s limbs. Moths, on the other hand, pollinate flowers during the late afternoon and night. The flowers pollinated by moths are pale or white and are flat, enabling the moths to land. One well-studied example of a moth-pollinated plant is the yucca plant, which is pollinated by the yucca moth. The shape of the flower and moth have adapted in such a way as to allow successful pollination. The moth deposits pollen on the sticky stigma for fertilization to occur later. The female moth also deposits eggs into the ovary. As the eggs develop into larvae, they obtain food from the flower and developing seeds. Thus, both the insect and flower benefit from each other in this symbiotic relationship. The corn earworm moth and Gaura plant have a similar relationship (Figure \(2\)).
Pollination by Bats
In the tropics and deserts, bats are often the pollinators of nocturnal flowers such as agave, guava, and morning glory. The flowers are usually large and white or pale-colored; thus, they can be distinguished from the dark surroundings at night. The flowers have a strong, fruity, or musky fragrance and produce large amounts of nectar. They are naturally large and wide-mouthed to accommodate the head of the bat. As the bats seek the nectar, their faces and heads become covered with pollen, which is then transferred to the next flower.
Pollination by Birds
Many species of small birds, such as the hummingbird (Figure \(3\)) and sun birds, are pollinators for plants such as orchids and other wildflowers. Flowers visited by birds are usually sturdy and are oriented in such a way as to allow the birds to stay near the flower without getting their wings entangled in the nearby flowers. The flower typically has a curved, tubular shape, which allows access for the bird’s beak. Brightly colored, odorless flowers that are open during the day are pollinated by birds. As a bird seeks energy-rich nectar, pollen is deposited on the bird’s head and neck and is then transferred to the next flower it visits. Botanists have been known to determine the range of extinct plants by collecting and identifying pollen from 200-year-old bird specimens from the same site.
Pollination by Wind
Most species of conifers, and many angiosperms, such as grasses, maples and oaks, are pollinated by wind. Pine cones are brown and unscented, while the flowers of wind-pollinated angiosperm species are usually green, small, may have small or no petals, and produce large amounts of pollen. Unlike the typical insect-pollinated flowers, flowers adapted to pollination by wind do not produce nectar or scent. In wind-pollinated species, the microsporangia hang out of the flower, and, as the wind blows, the lightweight pollen is carried with it (Figure \(4\)). The flowers usually emerge early in the spring, before the leaves, so that the leaves do not block the movement of the wind. The pollen is deposited on the exposed feathery stigma of the flower (Figure \(5\)).
Pollination by Water
Some weeds, such as Australian sea grass and pond weeds, are pollinated by water. The pollen floats on water, and when it comes into contact with the flower, it is deposited inside the flower.
Evolution Connection: Pollination by Deception
Orchids are highly valued flowers, with many rare varieties (Figure \(6\)). They grow in a range of specific habitats, mainly in the tropics of Asia, South America, and Central America. At least 25,000 species of orchids have been identified.
Flowers often attract pollinators with food rewards, in the form of nectar. However, some species of orchid are an exception to this standard: they have evolved different ways to attract the desired pollinators. They use a method known as food deception, in which bright colors and perfumes are offered, but no food. Anacamptis morio, commonly known as the green-winged orchid, bears bright purple flowers and emits a strong scent. The bumblebee, its main pollinator, is attracted to the flower because of the strong scent—which usually indicates food for a bee—and in the process, picks up the pollen to be transported to another flower.
Other orchids use sexual deception. Chiloglottis trapeziformis emits a compound that smells the same as the pheromone emitted by a female wasp to attract male wasps. The male wasp is attracted to the scent, lands on the orchid flower, and in the process, transfers pollen. Some orchids, like the Australian hammer orchid, use scent as well as visual trickery in yet another sexual deception strategy to attract wasps. The flower of this orchid mimics the appearance of a female wasp and emits a pheromone. The male wasp tries to mate with what appears to be a female wasp, and in the process, picks up pollen, which it then transfers to the next counterfeit mate.
Double Fertilization
After pollen is deposited on the stigma, it must germinate and grow through the style to reach the ovule. The microspores, or the pollen, contain two cells: the pollen tube cell and the generative cell. The pollen tube cell grows into a pollen tube through which the generative cell travels. The germination of the pollen tube requires water, oxygen, and certain chemical signals. As it travels through the style to reach the embryo sac, the pollen tube’s growth is supported by the tissues of the style. In the meantime, if the generative cell has not already split into two cells, it now divides to form two sperm cells. The pollen tube is guided by the chemicals secreted by the synergids present in the embryo sac, and it enters the ovule sac through the micropyle. Of the two sperm cells, one sperm fertilizes the egg cell, forming a diploid zygote; the other sperm fuses with the two polar nuclei, forming a triploid cell that develops into the endosperm. Together, these two fertilization events in angiosperms are known as double fertilization (Figure \(7\)). After fertilization is complete, no other sperm can enter. The fertilized ovule forms the seed, whereas the tissues of the ovary become the fruit, usually enveloping the seed.
After fertilization, the zygote divides to form two cells: the upper cell, or terminal cell, and the lower, or basal, cell. The division of the basal cell gives rise to the suspensor, which eventually makes connection with the maternal tissue. The suspensor provides a route for nutrition to be transported from the mother plant to the growing embryo. The terminal cell also divides, giving rise to a globular-shaped proembryo (Figure \(8\)a). In dicots (eudicots), the developing embryo has a heart shape, due to the presence of the two rudimentary cotyledons (Figure \(8\)b). In non-endospermic dicots, such as Capsella bursa, the endosperm develops initially, but is then digested, and the food reserves are moved into the two cotyledons. As the embryo and cotyledons enlarge, they run out of room inside the developing seed, and are forced to bend (Figure \(8\)c). Ultimately, the embryo and cotyledons fill the seed (Figure \(8\)d), and the seed is ready for dispersal. Embryonic development is suspended after some time, and growth is resumed only when the seed germinates. The developing seedling will rely on the food reserves stored in the cotyledons until the first set of leaves begin photosynthesis.
Development of the Seed
The mature ovule develops into the seed. A typical seed contains a seed coat, cotyledons, endosperm, and a single embryo (Figure \(9\)).
Art Connection
Which of the following statements is true?
1. Both monocots and dicots have an endosperm.
2. The radicle develops into the root.
3. The plumule is part of the epicotyl.
4. The endosperm is part of the embryo.
The storage of food reserves in angiosperm seeds differs between monocots and dicots. In monocots, such as corn and wheat, the single cotyledon is called a scutellum; the scutellum is connected directly to the embryo via vascular tissue (xylem and phloem). Food reserves are stored in the large endosperm. Upon germination, enzymes are secreted by the aleurone, a single layer of cells just inside the seed coat that surrounds the endosperm and embryo. The enzymes degrade the stored carbohydrates, proteins and lipids, the products of which are absorbed by the scutellum and transported via a vasculature strand to the developing embryo. Therefore, the scutellum can be seen to be an absorptive organ, not a storage organ.
The two cotyledons in the dicot seed also have vascular connections to the embryo. In endospermic dicots, the food reserves are stored in the endosperm. During germination, the two cotyledons therefore act as absorptive organs to take up the enzymatically released food reserves, much like in monocots (monocots, by definition, also have endospermic seeds). Tobacco (Nicotiana tabaccum), tomato (Solanum lycopersicum), and pepper (Capsicum annuum) are examples of endospermic dicots. In non-endospermic dicots, the triploid endosperm develops normally following double fertilization, but the endosperm food reserves are quickly remobilized and moved into the developing cotyledon for storage. The two halves of a peanut seed (Arachis hypogaea) and the split peas (Pisum sativum) of split pea soup are individual cotyledons loaded with food reserves.
The seed, along with the ovule, is protected by a seed coat that is formed from the integuments of the ovule sac. In dicots, the seed coat is further divided into an outer coat known as the testa and inner coat known as the tegmen.
The embryonic axis consists of three parts: the plumule, the radicle, and the hypocotyl. The portion of the embryo between the cotyledon attachment point and the radicle is known as the hypocotyl (hypocotyl means “below the cotyledons”). The embryonic axis terminates in a radicle (the embryonic root), which is the region from which the root will develop. In dicots, the hypocotyls extend above ground, giving rise to the stem of the plant. In monocots, the hypocotyl does not show above ground because monocots do not exhibit stem elongation. The part of the embryonic axis that projects above the cotyledons is known as the epicotyl. The plumule is composed of the epicotyl, young leaves, and the shoot apical meristem.
Upon germination in dicot seeds, the epicotyl is shaped like a hook with the plumule pointing downwards. This shape is called the plumule hook, and it persists as long as germination proceeds in the dark. Therefore, as the epicotyl pushes through the tough and abrasive soil, the plumule is protected from damage. Upon exposure to light, the hypocotyl hook straightens out, the young foliage leaves face the sun and expand, and the epicotyl continues to elongate. During this time, the radicle is also growing and producing the primary root. As it grows downward to form the tap root, lateral roots branch off to all sides, producing the typical dicot tap root system.
In monocot seeds (Figure \(10\)), the testa and tegmen of the seed coat are fused. As the seed germinates, the primary root emerges, protected by the root-tip covering: the coleorhiza. Next, the primary shoot emerges, protected by the coleoptile: the covering of the shoot tip. Upon exposure to light (i.e. when the plumule has exited the soil and the protective coleoptile is no longer needed), elongation of the coleoptile ceases and the leaves expand and unfold. At the other end of the embryonic axis, the primary root soon dies, while other, adventitious roots (roots that do not arise from the usual place – i.e. the root) emerge from the base of the stem. This gives the monocot a fibrous root system.
Seed Germination
Many mature seeds enter a period of inactivity, or extremely low metabolic activity: a process known as dormancy, which may last for months, years or even centuries. Dormancy helps keep seeds viable during unfavorable conditions. Upon a return to favorable conditions, seed germination takes place. Favorable conditions could be as diverse as moisture, light, cold, fire, or chemical treatments. After heavy rains, many new seedlings emerge. Forest fires also lead to the emergence of new seedlings. Some seeds require vernalization (cold treatment) before they can germinate. This guarantees that seeds produced by plants in temperate climates will not germinate until the spring. Plants growing in hot climates may have seeds that need a heat treatment in order to germinate, to avoid germination in the hot, dry summers. In many seeds, the presence of a thick seed coat retards the ability to germinate. Scarification, which includes mechanical or chemical processes to soften the seed coat, is often employed before germination. Presoaking in hot water, or passing through an acid environment, such as an animal’s digestive tract, may also be employed.
Depending on seed size, the time taken for a seedling to emerge may vary. Species with large seeds have enough food reserves to germinate deep below ground, and still extend their epicotyl all the way to the soil surface. Seeds of small-seeded species usually require light as a germination cue. This ensures the seeds only germinate at or near the soil surface (where the light is greatest). If they were to germinate too far underneath the surface, the developing seedling would not have enough food reserves to reach the sunlight.
Development of Fruit and Fruit Types
After fertilization, the ovary of the flower usually develops into the fruit. Fruits are usually associated with having a sweet taste; however, not all fruits are sweet. Botanically, the term “fruit” is used for a ripened ovary. In most cases, flowers in which fertilization has taken place will develop into fruits, and flowers in which fertilization has not taken place will not. Some fruits develop from the ovary and are known as true fruits, whereas others develop from other parts of the female gametophyte and are known as accessory fruits. The fruit encloses the seeds and the developing embryo, thereby providing it with protection. Fruits are of many types, depending on their origin and texture. The sweet tissue of the blackberry, the red flesh of the tomato, the shell of the peanut, and the hull of corn (the tough, thin part that gets stuck in your teeth when you eat popcorn) are all fruits. As the fruit matures, the seeds also mature.
Fruits may be classified as simple, aggregate, multiple, or accessory, depending on their origin (Figure \(11\)). If the fruit develops from a single carpel or fused carpels of a single ovary, it is known as a simple fruit, as seen in nuts and beans. An aggregate fruit is one that develops from more than one carpel, but all are in the same flower: the mature carpels fuse together to form the entire fruit, as seen in the raspberry. Multiple fruit develops from an inflorescence or a cluster of flowers. An example is the pineapple, where the flowers fuse together to form the fruit. Accessory fruits (sometimes called false fruits) are not derived from the ovary, but from another part of the flower, such as the receptacle (strawberry) or the hypanthium (apples and pears).
Fruits generally have three parts: the exocarp (the outermost skin or covering), the mesocarp (middle part of the fruit), and the endocarp (the inner part of the fruit). Together, all three are known as the pericarp. The mesocarp is usually the fleshy, edible part of the fruit; however, in some fruits, such as the almond, the endocarp is the edible part. In many fruits, two or all three of the layers are fused, and are indistinguishable at maturity. Fruits can be dry or fleshy. Furthermore, fruits can be divided into dehiscent or indehiscent types. Dehiscent fruits, such as peas, readily release their seeds, while indehiscent fruits, like peaches, rely on decay to release their seeds.
Fruit and Seed Dispersal
The fruit has a single purpose: seed dispersal. Seeds contained within fruits need to be dispersed far from the mother plant, so they may find favorable and less competitive conditions in which to germinate and grow.
Some fruit have built-in mechanisms so they can disperse by themselves, whereas others require the help of agents like wind, water, and animals (Figure \(12\)). Modifications in seed structure, composition, and size help in dispersal. Wind-dispersed fruit are lightweight and may have wing-like appendages that allow them to be carried by the wind. Some have a parachute-like structure to keep them afloat. Some fruits—for example, the dandelion—have hairy, weightless structures that are suited to dispersal by wind.
Seeds dispersed by water are contained in light and buoyant fruit, giving them the ability to float. Coconuts are well known for their ability to float on water to reach land where they can germinate. Similarly, willow and silver birches produce lightweight fruit that can float on water.
Animals and birds eat fruits, and the seeds that are not digested are excreted in their droppings some distance away. Some animals, like squirrels, bury seed-containing fruits for later use; if the squirrel does not find its stash of fruit, and if conditions are favorable, the seeds germinate. Some fruits, like the cocklebur, have hooks or sticky structures that stick to an animal's coat and are then transported to another place. Humans also play a big role in dispersing seeds when they carry fruits to new places and throw away the inedible part that contains the seeds.
All of the above mechanisms allow for seeds to be dispersed through space, much like an animal’s offspring can move to a new location. Seed dormancy, which was described earlier, allows plants to disperse their progeny through time: something animals cannot do. Dormant seeds can wait months, years, or even decades for the proper conditions for germination and propagation of the species.
Summary
For fertilization to occur in angiosperms, pollen has to be transferred to the stigma of a flower: a process known as pollination. Gymnosperm pollination involves the transfer of pollen from a male cone to a female cone. When the pollen of the flower is transferred to the stigma of the same flower, it is called self-pollination. Cross-pollination occurs when pollen is transferred from one flower to another flower on the same plant, or another plant. Cross-pollination requires pollinating agents such as water, wind, or animals, and increases genetic diversity. After the pollen lands on the stigma, the tube cell gives rise to the pollen tube, through which the generative nucleus migrates. The pollen tube gains entry through the micropyle on the ovule sac. The generative cell divides to form two sperm cells: one fuses with the egg to form the diploid zygote, and the other fuses with the polar nuclei to form the endosperm, which is triploid in nature. This is known as double fertilization. After fertilization, the zygote divides to form the embryo and the fertilized ovule forms the seed. The walls of the ovary form the fruit in which the seeds develop. The seed, when mature, will germinate under favorable conditions and give rise to the diploid sporophyte.
Art Connections
Figure \(9\): What is the function of the cotyledon?
1. It develops into the root.
2. It provides nutrition for the embryo.
3. It forms the embryo.
4. It protects the embryo.
Answer
B
Glossary
accessory fruit
fruit derived from tissues other than the ovary
aggregate fruit
fruit that develops from multiple carpels in the same flower
aleurone
single layer of cells just inside the seed coat that secretes enzymes upon germination
coleoptile
covering of the shoot tip, found in germinating monocot seeds
coleorhiza
covering of the root tip, found in germinating monocot seeds
cotyledon
fleshy part of seed that provides nutrition to the seed
cross-pollination
transfer of pollen from the anther of one flower to the stigma of a different flower
dormancy
period of no growth and very slow metabolic processes
double fertilization
two fertilization events in angiosperms; one sperm fuses with the egg, forming the zygote, whereas the other sperm fuses with the polar nuclei, forming endosperm
endocarp
innermost part of fruit
endosperm
triploid structure resulting from fusion of a sperm with polar nuclei, which serves as a nutritive tissue for embryo
endospermic dicot
dicot that stores food reserves in the endosperm
exocarp
outermost covering of a fruit
epicotyl
embryonic shoot above the cotyledons
gravitropism
response of a plant growth in the same direction as gravity
hypocotyl
embryonic axis above the cotyledons
mesocarp
middle part of a fruit
multiple fruit
fruit that develops from multiple flowers on an inflorescence
nectar guide
pigment pattern on a flower that guides an insect to the nectaries
non-endospermic dicot
dicot that stores food reserves in the developing cotyledon
pericarp
collective term describing the exocarp, mesocarp, and endocarp; the structure that encloses the seed and is a part of the fruit
plumule
shoot that develops from the germinating seed
pollination
transfer of pollen to the stigma
radicle
original root that develops from the germinating seed
scarification
mechanical or chemical processes to soften the seed coat
scutellum
type of cotyledon found in monocots, as in grass seeds
self-pollination
transfer of pollen from the anther to the stigma of same flower
simple fruit
fruit that develops from a single carpel or fused carpels
suspensor
part of the growing embryo that makes connection with the maternal tissues
tegmen
inner layer of the seed coat
testa
outer layer of the seed coat
vernalization
exposure to cold required by some seeds before they can germinate | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/32%3A_Plant_Reproduction/32.2%3A_Pollination_and_Fertilization.txt |
Skills to Develop
• Compare the mechanisms and methods of natural and artificial asexual reproduction
• Describe the advantages and disadvantages of natural and artificial asexual reproduction
• Discuss plant life spans
Many plants are able to propagate themselves using asexual reproduction. This method does not require the investment required to produce a flower, attract pollinators, or find a means of seed dispersal. Asexual reproduction produces plants that are genetically identical to the parent plant because no mixing of male and female gametes takes place. Traditionally, these plants survive well under stable environmental conditions when compared with plants produced from sexual reproduction because they carry genes identical to those of their parents.
Many different types of roots exhibit asexual reproduction Figure \(1\). The corm is used by gladiolus and garlic. Bulbs, such as a scaly bulb in lilies and a tunicate bulb in daffodils, are other common examples. A potato is a stem tuber, while parsnip propagates from a taproot. Ginger and iris produce rhizomes, while ivy uses an adventitious root (a root arising from a plant part other than the main or primary root), and the strawberry plant has a stolon, which is also called a runner.
Some plants can produce seeds without fertilization. Either the ovule or part of the ovary, which is diploid in nature, gives rise to a new seed. This method of reproduction is known as apomixis.
An advantage of asexual reproduction is that the resulting plant will reach maturity faster. Since the new plant is arising from an adult plant or plant parts, it will also be sturdier than a seedling. Asexual reproduction can take place by natural or artificial (assisted by humans) means.
Natural Methods of Asexual Reproduction
Natural methods of asexual reproduction include strategies that plants have developed to self-propagate. Many plants—like ginger, onion, gladioli, and dahlia—continue to grow from buds that are present on the surface of the stem. In some plants, such as the sweet potato, adventitious roots or runners can give rise to new plants Figure \(2\). In Bryophyllum and kalanchoe, the leaves have small buds on their margins. When these are detached from the plant, they grow into independent plants; or, they may start growing into independent plants if the leaf touches the soil. Some plants can be propagated through cuttings alone.
Artificial Methods of Asexual Reproduction
These methods are frequently employed to give rise to new, and sometimes novel, plants. They include grafting, cutting, layering, and micropropagation.
Grafting
Grafting has long been used to produce novel varieties of roses, citrus species, and other plants. In grafting, two plant species are used; part of the stem of the desirable plant is grafted onto a rooted plant called the stock. The part that is grafted or attached is called the scion. Both are cut at an oblique angle (any angle other than a right angle), placed in close contact with each other, and are then held together Figure \(3\). Matching up these two surfaces as closely as possible is extremely important because these will be holding the plant together. The vascular systems of the two plants grow and fuse, forming a graft. After a period of time, the scion starts producing shoots, and eventually starts bearing flowers and fruits. Grafting is widely used in viticulture (grape growing) and the citrus industry. Scions capable of producing a particular fruit variety are grated onto root stock with specific resistance to disease.
Cutting
Plants such as coleus and money plant are propagated through stem cuttings, where a portion of the stem containing nodes and internodes is placed in moist soil and allowed to root. In some species, stems can start producing a root even when placed only in water. For example, leaves of the African violet will root if kept in water undisturbed for several weeks.
Layering
Layering is a method in which a stem attached to the plant is bent and covered with soil. Young stems that can be bent easily without any injury are preferred. Jasmine and bougainvillea (paper flower) can be propagated this way Figure \(4\). In some plants, a modified form of layering known as air layering is employed. A portion of the bark or outermost covering of the stem is removed and covered with moss, which is then taped. Some gardeners also apply rooting hormone. After some time, roots will appear, and this portion of the plant can be removed and transplanted into a separate pot.
Micropropagation
Micropropagation (also called plant tissue culture) is a method of propagating a large number of plants from a single plant in a short time under laboratory conditions Figure \(5\). This method allows propagation of rare, endangered species that may be difficult to grow under natural conditions, are economically important, or are in demand as disease-free plants.
To start plant tissue culture, a part of the plant such as a stem, leaf, embryo, anther, or seed can be used. The plant material is thoroughly sterilized using a combination of chemical treatments standardized for that species. Under sterile conditions, the plant material is placed on a plant tissue culture medium that contains all the minerals, vitamins, and hormones required by the plant. The plant part often gives rise to an undifferentiated mass known as callus, from which individual plantlets begin to grow after a period of time. These can be separated and are first grown under greenhouse conditions before they are moved to field conditions.
Plant Life Spans
The length of time from the beginning of development to the death of a plant is called its life span. The life cycle, on the other hand, is the sequence of stages a plant goes through from seed germination to seed production of the mature plant. Some plants, such as annuals, only need a few weeks to grow, produce seeds and die. Other plants, such as the bristlecone pine, live for thousands of years. Some bristlecone pines have a documented age of 4,500 years Figure \(6\). Even as some parts of a plant, such as regions containing meristematic tissue—the area of active plant growth consisting of undifferentiated cells capable of cell division—continue to grow, some parts undergo programmed cell death (apoptosis). The cork found on stems, and the water-conducting tissue of the xylem, for example, are composed of dead cells.
Plant species that complete their lifecycle in one season are known as annuals, an example of which is Arabidopsis, or mouse-ear cress. Biennials such as carrots complete their lifecycle in two seasons. In a biennial’s first season, the plant has a vegetative phase, whereas in the next season, it completes its reproductive phase. Commercial growers harvest the carrot roots after the first year of growth, and do not allow the plants to flower. Perennials, such as the magnolia, complete their lifecycle in two years or more.
In another classification based on flowering frequency, monocarpic plants flower only once in their lifetime; examples include bamboo and yucca. During the vegetative period of their life cycle (which may be as long as 120 years in some bamboo species), these plants may reproduce asexually and accumulate a great deal of food material that will be required during their once-in-a-lifetime flowering and setting of seed after fertilization. Soon after flowering, these plants die. Polycarpic plants form flowers many times during their lifetime. Fruit trees, such as apple and orange trees, are polycarpic; they flower every year. Other polycarpic species, such as perennials, flower several times during their life span, but not each year. By this means, the plant does not require all its nutrients to be channelled towards flowering each year.
As is the case with all living organisms, genetics and environmental conditions have a role to play in determining how long a plant will live. Susceptibility to disease, changing environmental conditions, drought, cold, and competition for nutrients are some of the factors that determine the survival of a plant. Plants continue to grow, despite the presence of dead tissue such as cork. Individual parts of plants, such as flowers and leaves, have different rates of survival. In many trees, the older leaves turn yellow and eventually fall from the tree. Leaf fall is triggered by factors such as a decrease in photosynthetic efficiency, due to shading by upper leaves, or oxidative damage incurred as a result of photosynthetic reactions. The components of the part to be shed are recycled by the plant for use in other processes, such as development of seed and storage. This process is known as nutrient recycling.
The aging of a plant and all the associated processes is known as senescence, which is marked by several complex biochemical changes. One of the characteristics of senescence is the breakdown of chloroplasts, which is characterized by the yellowing of leaves. The chloroplasts contain components of photosynthetic machinery such as membranes and proteins. Chloroplasts also contain DNA. The proteins, lipids, and nucleic acids are broken down by specific enzymes into smaller molecules and salvaged by the plant to support the growth of other plant tissues.
The complex pathways of nutrient recycling within a plant are not well understood. Hormones are known to play a role in senescence. Applications of cytokinins and ethylene delay or prevent senescence; in contrast, abscissic acid causes premature onset of senescence.
Summary
Many plants reproduce asexually as well as sexually. In asexual reproduction, part of the parent plant is used to generate a new plant. Grafting, layering, and micropropagation are some methods used for artificial asexual reproduction. The new plant is genetically identical to the parent plant from which the stock has been taken. Asexually reproducing plants thrive well in stable environments.
Plants have different life spans, dependent on species, genotype, and environmental conditions. Parts of the plant, such as regions containing meristematic tissue, continue to grow, while other parts experience programmed cell death. Leaves that are no longer photosynthetically active are shed from the plant as part of senescence, and the nutrients from these leaves are recycled by the plant. Other factors, including the presence of hormones, are known to play a role in delaying senescence.
Glossary
apomixis
process by which seeds are produced without fertilization of sperm and egg
cutting
method of asexual reproduction where a portion of the stem contains notes and internodes is placed in moist soil and allowed to root
grafting
method of asexual reproduction where the stem from one plant species is spliced to a different plant
layering
method of propagating plants by bending a stem under the soil
micropropagation
propagation of desirable plants from a plant part; carried out in a laboratory
monocarpic
plants that flower once in their lifetime
polycarpic
plants that flower several times in their lifetime
scion
the part of a plant that is grafted onto the root stock of another plant
senescence
process that describes aging in plant tissues | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/32%3A_Plant_Reproduction/32.3%3A_Asexual_Reproduction.txt |
32.1: Reproductive Development and Structure
Sexual reproduction takes place with slight variations in different groups of plants. Plants have two distinct stages in their lifecycle: the gametophyte stage and the sporophyte stage. The haploid gametophyte produces the male and female gametes by mitosis in distinct multicellular structures. Fusion of the male and females gametes forms the diploid zygote, which develops into the sporophyte.
Review Questions
In a plant’s male reproductive organs, development of pollen takes place in a structure known as the ________.
1. stamen
2. microsporangium
3. anther
4. tapetum
Answer
B
The stamen consists of a long stalk called the filament that supports the ________.
1. stigma
2. sepal
3. style
4. anther
Answer
D
The ________ are collectively called the calyx.
1. sepals
2. petals
3. tepals
4. stamens
Answer
A
The pollen lands on which part of the flower?
1. stigma
2. style
3. ovule
4. integument
Answer
A
Free Response
Describe the reproductive organs inside a flower.
Answer
Inside the flower are the reproductive organs of the plant. The stamen is the male reproductive organ. Pollen is produced in the stamen. The carpel is the female reproductive organ. The ovary is the swollen base of the carpel where ovules are found. Not all flowers have every one of the four parts.
Describe the two-stage lifecycle of plants: the gametophyte stage and the sporophyte stage.
Answer
Plants have two distinct phases in their lifecycle: the gametophyte stage and the sporophyte stage. In the gametophyte stage, when reproductive cells undergo meiosis and produce haploid cells called spores, the gametophyte stage begins. Spores divide by cell division to form plant structures of an entirely new plant. The cells in these structures or plants are haploid. Some of these cells undergo cell division and form sex cells. Fertilization, the joining of haploid sex cells, begins the sporophyte stage. Cells formed in this stage have the diploid number of chromosomes. Meiosis in some of these cells forms spores, and the cycle begins again: a process known as alternation of generations.
Describe the four main parts, or whorls, of a flower.
Answer
A typical flower has four main parts, or whorls: the calyx, corolla, androecium, and gynoecium. The outermost whorl of the flower has green, leafy structures known as sepals, which are collectively called the calyx. It helps to protect the unopened bud. The second whorl is made up of brightly colored petals that are known collectively as the corolla. The third whorl is the male reproductive structure known as the androecium. The androecium has stamens, which have anthers on a stalk or filament. Pollen grains are borne on the anthers. The gynoecium is the female reproductive structure. The carpel is the individual structure of the gynoecium and has a stigma, the stalk or style, and the ovary.
Discuss the differences between a complete flower and an incomplete flower.
Answer
If all four whorls of a flower are present, it is a complete flower. If any of the four parts is missing, it is known as incomplete. Flowers that contain both an androecium and gynoecium are called androgynous or hermaphrodites. Those that contain only an androecium are known as staminate flowers, and those that have only carpels are known as carpellate. If both male and female flowers are borne on the same plant, it is called monoecious, while plants with male and female flowers on separate plants are termed dioecious.
32.2: Pollination and Fertilization
Pollination takes two forms: self-pollination and cross-pollination. Self-pollination occurs when the pollen from the anther is deposited on the stigma of the same flower, or another flower on the same plant. Cross-pollination is the transfer of pollen from the anther of one flower to the stigma of another flower on a different individual of the same species. Self-pollination occurs in flowers where the stamen and carpel mature at the same time.
Review Questions
After double fertilization, a zygote and ________ form.
1. an ovule
2. endosperm
3. a cotyledon
4. a suspensor
Answer
B
The fertilized ovule gives rise to the ________.
1. fruit
2. seed
3. endosperm
4. embryo
Answer
B
What is the term for a fruit that develops from tissues other than the ovary?
1. simple fruit
2. aggregate fruit
3. multiple fruit
4. accessory fruit
Answer
D
The ________ is the outermost covering of a fruit.
1. endocarp
2. pericarp
3. exocarp
4. mesocarp
Answer
C
Free Response
Why do some seeds undergo a period of dormancy, and how do they break dormancy?
Answer
Many seeds enter a period of inactivity or extremely low metabolic activity, a process known as dormancy. Dormancy allows seeds to tide over unfavorable conditions and germinate on return to favorable conditions. Favorable conditions could be as diverse as moisture, light, cold, fire, or chemical treatments. After heavy rains, many new seedlings emerge. Forest fires also lead to the emergence of new seedlings.
Discuss some ways in which fruit seeds are dispersed.
Answer
Some fruits have built-in mechanisms that allow them to disperse seeds by themselves, but others require the assistance of agents like wind, water, and animals. Fruit that are dispersed by the wind are light in weight and often have wing-like appendages that allow them to be carried by the wind; other have structures resembling a parachute that keep them afloat in the wind. Some fruits, such as those of dandelions, have hairy, weightless structures that allow them to float in the wind. Fruits dispersed by water are light and buoyant, giving them the ability to float; coconuts are one example. Animals and birds eat fruits and disperse their seeds by leaving droppings at distant locations. Other animals bury fruit that may later germinate. Some fruits stick to animals’ bodies and are carried to new locations. People also contribute to seed dispersal when they carry fruits to new places.
32.3: Asexual Reproduction
Many plants are able to propagate themselves using asexual reproduction. This method does not require the investment required to produce a flower, attract pollinators, or find a means of seed dispersal. Asexual reproduction produces plants that are genetically identical to the parent plant because no mixing of male and female gametes takes place. Traditionally, these plants survive well under stable environmental conditions when compared with plants produced from sexual reproduction.
Review Questions
________ is a useful method of asexual reproduction for propagating hard-to-root plants.
1. grafting
2. layering
3. cuttings
4. budding
Answer
A
Which of the following is an advantage of asexual reproduction?
1. Cuttings taken from an adult plant show increased resistance to diseases.
2. Grafted plants can more successfully endure drought.
3. When cuttings or buds are taken from an adult plant or plant parts, the resulting plant will grow into an adult faster than a seedling.
4. Asexual reproduction takes advantage of a more diverse gene pool.
Answer
C
Plants that flower once in their lifetime are known as ________.
1. monoecious
2. dioecious
3. polycarpic
4. monocarpic
Answer
D
Plant species that complete their lifecycle in one season are known as ________.
1. biennials
2. perennials
3. annuals
4. polycarpic
Answer
C
Free Response
What are some advantages of asexual reproduction in plants?
Answer
Asexual reproduction does not require the expenditure of the plant’s resources and energy that would be involved in producing a flower, attracting pollinators, or dispersing seeds. Asexual reproduction results in plants that are genetically identical to the parent plant, since there is no mixing of male and female gametes, resulting in better survival. The cuttings or buds taken from an adult plant produce progeny that mature faster and are sturdier than a seedling grown from a seed.
Describe natural and artificial methods of asexual reproduction in plants.
Answer
Asexual reproduction in plants can take place by natural methods or artificial methods. Natural methods include strategies used by the plant to propagate itself. Artificial methods include grafting, cutting, layering, and micropropagation.
Discuss the life cycles of various plants.
Answer
Plant species that complete their life cycle in one season are known as annuals. Biennials complete their life cycle in two seasons. In the first season, the plant has a vegetative phase, whereas in the next season, it completes its reproductive phase. Perennials, such as the magnolia, complete their life cycle in two years or more.
How are plants classified on the basis of flowering frequency?
Answer
Monocarpic plants flower only once during their lifetime. During the vegetative period of their lifecycle, these plants accumulate a great deal of food material that will be required during their once-in-a-lifetime flowering and setting of seed after fertilization. Soon after flowering, these plants die. Polycarpic plants flower several times during their life span; therefore, not all nutrients are channelled towards flowering. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/6%3A_Plant_Structure_and_Function/32%3A_Plant_Reproduction/32.E%3A_Plant_Reproduction_%28Exercises%29.txt |
The structures of animals consist of primary tissues that make up more complex organs and organ systems. Homeostasis allows an animal to maintain a balance between its internal and external environments.
• 33.0: Prelude to The Animal Body
The structures of animals consist of primary tissues that make up more complex organs and organ systems. Homeostasis allows an animal to maintain a balance between its internal and external environments.
• 33.1: Animal Form and Function
Animals vary in form and function. From a sponge to a worm to a goat, an organism has a distinct body plan that limits its size and shape. Animals’ bodies are also designed to interact with their environments, whether in the deep sea, a rainforest canopy, or the desert. Therefore, a large amount of information about the structure of an organism's body (anatomy) and the function of its cells, tissues and organs (physiology) can be learned by studying that organism's environment.
• 33.2: Animal Primary Tissues
The tissues of multicellular, complex animals are four primary types: epithelial, connective, muscle, and nervous. Recall that tissues are groups of similar cells group of similar cells carrying out related functions. These tissues combine to form organs—like the skin or kidney—that have specific, specialized functions within the body. Organs are organized into organ systems to perform functions.
• 33.3: Homeostasis
Animal organs and organ systems constantly adjust to internal and external changes through a process called homeostasis (“steady state”). These changes might be in the level of glucose or calcium in blood or in external temperatures. Homeostasis means to maintain dynamic equilibrium in the body. It is dynamic because it is constantly adjusting to the changes that the body’s systems encounter. It is equilibrium because body functions are kept within specific ranges.
• 33.E: The Animal Body - Basic Form and Function (Exercises)
Thumbnail: Elephant. (CC BY 2.0/cropped from original; Caitlin via Flickr).
33: The Animal Body - Basic Form and Function
The arctic fox is an example of a complex animal that has adapted to its environment and illustrates the relationships between an animal’s form and function. The structures of animals consist of primary tissues that make up more complex organs and organ systems. Homeostasis allows an animal to maintain a balance between its internal and external environments.
33.1: Animal Form and Function
Skills to Develop
• Describe the various types of body plans that occur in animals
• Describe limits on animal size and shape
• Relate bioenergetics to body size, levels of activity, and the environment
Animals vary in form and function. From a sponge to a worm to a goat, an organism has a distinct body plan that limits its size and shape. Animals’ bodies are also designed to interact with their environments, whether in the deep sea, a rainforest canopy, or the desert. Therefore, a large amount of information about the structure of an organism's body (anatomy) and the function of its cells, tissues and organs (physiology) can be learned by studying that organism's environment.
Body Plans
Animal body plans follow set patterns related to symmetry. They are asymmetrical, radial, or bilateral in form as illustrated in Figure \(1\). Asymmetrical animals are animals with no pattern or symmetry; an example of an asymmetrical animal is a sponge. Radial symmetry, as illustrated in Figure \(1\), describes when an animal has an up-and-down orientation: any plane cut along its longitudinal axis through the organism produces equal halves, but not a definite right or left side. This plan is found mostly in aquatic animals, especially organisms that attach themselves to a base, like a rock or a boat, and extract their food from the surrounding water as it flows around the organism. Bilateral symmetry is illustrated in the same figure by a goat. The goat also has an upper and lower component to it, but a plane cut from front to back separates the animal into definite right and left sides. Additional terms used when describing positions in the body are anterior (front), posterior (rear), dorsal (toward the back), and ventral (toward the stomach). Bilateral symmetry is found in both land-based and aquatic animals; it enables a high level of mobility.
Limits on Animal Size and Shape
Animals with bilateral symmetry that live in water tend to have a fusiform shape: this is a tubular shaped body that is tapered at both ends. This shape decreases the drag on the body as it moves through water and allows the animal to swim at high speeds. The table below lists the maximum speed of various animals. Certain types of sharks can swim at fifty kilometers an hour and some dolphins at 32 to 40 kilometers per hour. Land animals frequently travel faster, although the tortoise and snail are significantly slower than cheetahs. Another difference in the adaptations of aquatic and land-dwelling organisms is that aquatic organisms are constrained in shape by the forces of drag in the water since water has higher viscosity than air. On the other hand, land-dwelling organisms are constrained mainly by gravity, and drag is relatively unimportant. For example, most adaptations in birds are for gravity not for drag.
Table \(1\): Maximum Speed of Assorted Land and Marine Animals
Animal Speed (kmh) Speed (mph)
Cheetah 113 70
Quarter horse 77 48
Fox 68 42
Shortfin mako shark 50 31
Domestic house cat 48 30
Human 45 28
Dolphin 32–40 20–25
Mouse 13 8
Snail 0.05 0.03
Most animals have an exoskeleton, including insects, spiders, scorpions, horseshoe crabs, centipedes, and crustaceans. Scientists estimate that, of insects alone, there are over 30 million species on our planet. The exoskeleton is a hard covering or shell that provides benefits to the animal, such as protection against damage from predators and from water loss (for land animals); it also provides for the attachments of muscles.
As the tough and resistant outer cover of an arthropod, the exoskeleton may be constructed of a tough polymer such as chitin and is often biomineralized with materials such as calcium carbonate. This is fused to the animal’s epidermis. Ingrowths of the exoskeleton, called apodemes, function as attachment sites for muscles, similar to tendons in more advanced animals (Figure \(2\)). In order to grow, the animal must first synthesize a new exoskeleton underneath the old one and then shed or molt the original covering. This limits the animal’s ability to grow continually, and may limit the individual’s ability to mature if molting does not occur at the proper time. The thickness of the exoskeleton must be increased significantly to accommodate any increase in weight. It is estimated that a doubling of body size increases body weight by a factor of eight. The increasing thickness of the chitin necessary to support this weight limits most animals with an exoskeleton to a relatively small size. The same principles apply to endoskeletons, but they are more efficient because muscles are attached on the outside, making it easier to compensate for increased mass.
An animal with an endoskeleton has its size determined by the amount of skeletal system it needs in order to support the other tissues and the amount of muscle it needs for movement. As the body size increases, both bone and muscle mass increase. The speed achievable by the animal is a balance between its overall size and the bone and muscle that provide support and movement.
Limiting Effects of Diffusion on Size and Development
The exchange of nutrients and wastes between a cell and its watery environment occurs through the process of diffusion. All living cells are bathed in liquid, whether they are in a single-celled organism or a multicellular one. Diffusion is effective over a specific distance and limits the size that an individual cell can attain. If a cell is a single-celled microorganism, such as an amoeba, it can satisfy all of its nutrient and waste needs through diffusion. If the cell is too large, then diffusion is ineffective and the center of the cell does not receive adequate nutrients nor is it able to effectively dispel its waste.
An important concept in understanding how efficient diffusion is as a means of transport is the surface to volume ratio. Recall that any three-dimensional object has a surface area and volume; the ratio of these two quantities is the surface-to-volume ratio. Consider a cell shaped like a perfect sphere: it has a surface area of 4πr2, and a volume of (4/3)πr3. The surface-to-volume ratio of a sphere is 3/r; as the cell gets bigger, its surface to volume ratio decreases, making diffusion less efficient. The larger the size of the sphere, or animal, the less surface area for diffusion it possesses.
The solution to producing larger organisms is for them to become multicellular. Specialization occurs in complex organisms, allowing cells to become more efficient at doing fewer tasks. For example, circulatory systems bring nutrients and remove waste, while respiratory systems provide oxygen for the cells and remove carbon dioxide from them. Other organ systems have developed further specialization of cells and tissues and efficiently control body functions. Moreover, surface-to-volume ratio applies to other areas of animal development, such as the relationship between muscle mass and cross-sectional surface area in supporting skeletons, and in the relationship between muscle mass and the generation of dissipation of heat.
Animal Bioenergetics
All animals must obtain their energy from food they ingest or absorb. These nutrients are converted to adenosine triphosphate (ATP) for short-term storage and use by all cells. Some animals store energy for slightly longer times as glycogen, and others store energy for much longer times in the form of triglycerides housed in specialized adipose tissues. No energy system is one hundred percent efficient, and an animal’s metabolism produces waste energy in the form of heat. If an animal can conserve that heat and maintain a relatively constant body temperature, it is classified as a warm-blooded animal and called an endotherm. The insulation used to conserve the body heat comes in the forms of fur, fat, or feathers. The absence of insulation in ectothermic animals increases their dependence on the environment for body heat.
The amount of energy expended by an animal over a specific time is called its metabolic rate. The rate is measured variously in joules, calories, or kilocalories (1000 calories). Carbohydrates and proteins contain about 4.5 to 5 kcal/g, and fat contains about 9 kcal/g. Metabolic rate is estimated as the basal metabolic rate (BMR) in endothermic animals at rest and as the standard metabolic rate (SMR) in ectotherms. Human males have a BMR of 1600 to 1800 kcal/day, and human females have a BMR of 1300 to 1500 kcal/day. Even with insulation, endothermal animals require extensive amounts of energy to maintain a constant body temperature. An ectotherm such as an alligator has an SMR of 60 kcal/day.
Energy Requirements Related to Body Size
Smaller endothermic animals have a greater surface area for their mass than larger ones (Figure \(3\)). Therefore, smaller animals lose heat at a faster rate than larger animals and require more energy to maintain a constant internal temperature. This results in a smaller endothermic animal having a higher BMR, per body weight, than a larger endothermic animal.
Energy Requirements Related to Levels of Activity
The more active an animal is, the more energy is needed to maintain that activity, and the higher its BMR or SMR. The average daily rate of energy consumption is about two to four times an animal’s BMR or SMR. Humans are more sedentary than most animals and have an average daily rate of only 1.5 times the BMR. The diet of an endothermic animal is determined by its BMR. For example: the type of grasses, leaves, or shrubs that an herbivore eats affects the number of calories that it takes in. The relative caloric content of herbivore foods, in descending order, is tall grasses > legumes > short grasses > forbs (any broad-leaved plant, not a grass) > subshrubs > annuals/biennials.
Energy Requirements Related to Environment
Animals adapt to extremes of temperature or food availability through torpor. Torpor is a process that leads to a decrease in activity and metabolism and allows animals to survive adverse conditions. Torpor can be used by animals for long periods, such as entering a state of hibernation during the winter months, in which case it enables them to maintain a reduced body temperature. During hibernation, ground squirrels can achieve an abdominal temperature of 0° C (32° F), while a bear’s internal temperature is maintained higher at about 37° C (99° F).
If torpor occurs during the summer months with high temperatures and little water, it is called estivation. Some desert animals use this to survive the harshest months of the year. Torpor can occur on a daily basis; this is seen in bats and hummingbirds. While endothermy is limited in smaller animals by surface to volume ratio, some organisms can be smaller and still be endotherms because they employ daily torpor during the part of the day that is coldest. This allows them to conserve energy during the colder parts of the day, when they consume more energy to maintain their body temperature.
Animal Body Planes and Cavities
A standing vertebrate animal can be divided by several planes. A sagittal plane divides the body into right and left portions. A midsagittal plane divides the body exactly in the middle, making two equal right and left halves. A frontal plane (also called a coronal plane) separates the front from the back. A transverse plane (or, horizontal plane) divides the animal into upper and lower portions. This is sometimes called a cross section, and, if the transverse cut is at an angle, it is called an oblique plane. Figure \(4\) illustrates these planes on a goat (a four-legged animal) and a human being.
Vertebrate animals have a number of defined body cavities, as illustrated in Figure \(5\). Two of these are major cavities that contain smaller cavities within them. The dorsal cavity contains the cranial and the vertebral (or spinal) cavities. The ventral cavity contains the thoracic cavity, which in turn contains the pleural cavity around the lungs and the pericardial cavity, which surrounds the heart. The ventral cavity also contains the abdominopelvic cavity, which can be separated into the abdominal and the pelvic cavities.
Career Connections: Physical Anthropologist
Physical anthropologists study the adaption, variability, and evolution of human beings, plus their living and fossil relatives. They can work in a variety of settings, although most will have an academic appointment at a university, usually in an anthropology department or a biology, genetics, or zoology department.
Non-academic positions are available in the automotive and aerospace industries where the focus is on human size, shape, and anatomy. Research by these professionals might range from studies of how the human body reacts to car crashes to exploring how to make seats more comfortable. Other non-academic positions can be obtained in museums of natural history, anthropology, archaeology, or science and technology. These positions involve educating students from grade school through graduate school. Physical anthropologists serve as education coordinators, collection managers, writers for museum publications, and as administrators. Zoos employ these professionals, especially if they have an expertise in primate biology; they work in collection management and captive breeding programs for endangered species. Forensic science utilizes physical anthropology expertise in identifying human and animal remains, assisting in determining the cause of death, and for expert testimony in trials.
Summary
Animal bodies come in a variety of sizes and shapes. Limits on animal size and shape include impacts to their movement. Diffusion affects their size and development. Bioenergetics describes how animals use and obtain energy in relation to their body size, activity level, and environment.
Glossary
apodeme
ingrowth of an animal’s exoskeleton that functions as an attachment site for muscles
asymmetrical
describes animals with no axis of symmetry in their body pattern
basal metabolic rate (BMR)
metabolic rate at rest in endothermic animals
dorsal cavity
body cavity on the posterior or back portion of an animal; includes the cranial and vertebral cavities
ectotherm
animal incapable of maintaining a relatively constant internal body temperature
endotherm
animal capable of maintaining a relatively constant internal body temperature
estivation
torpor in response to extremely high temperatures and low water availability
frontal (coronal) plane
plane cutting through an animal separating the individual into front and back portions
fusiform
animal body shape that is tubular and tapered at both ends
hibernation
torpor over a long period of time, such as a winter
midsagittal plane
plane cutting through an animal separating the individual into even right and left sides
sagittal plane
plane cutting through an animal separating the individual into right and left sides
standard metabolic rate (SMR)
metabolic rate at rest in ectothermic animals
torpor
decrease in activity and metabolism that allows an animal to survive adverse conditions
transverse (horizontal) plane
plane cutting through an animal separating the individual into upper and lower portions
ventral cavity
body cavity on the anterior or front portion of an animal that includes the thoracic cavities and the abdominopelvic cavities | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/33%3A_The_Animal_Body_-_Basic_Form_and_Function/33.0%3A_Prelude_to_The_Animal_Body.txt |
Skills to Develop
• Describe epithelial tissues
• Discuss the different types of connective tissues in animals
• Describe three types of muscle tissues
• Describe nervous tissue
The tissues of multicellular, complex animals are four primary types: epithelial, connective, muscle, and nervous. Recall that tissues are groups of similar cells group of similar cells carrying out related functions. These tissues combine to form organs—like the skin or kidney—that have specific, specialized functions within the body. Organs are organized into organ systems to perform functions; examples include the circulatory system, which consists of the heart and blood vessels, and the digestive system, consisting of several organs, including the stomach, intestines, liver, and pancreas. Organ systems come together to create an entire organism.
Epithelial Tissues
Epithelial tissues cover the outside of organs and structures in the body and line the lumens of organs in a single layer or multiple layers of cells. The types of epithelia are classified by the shapes of cells present and the number of layers of cells. Epithelia composed of a single layer of cells is called simple epithelia; epithelial tissue composed of multiple layers is called stratified epithelia. The table summarizes the different types of epithelial tissues.
Table \(1\): Different Types of Epithelial Tissues
Cell shape Description Location
squamous flat, irregular round shape simple: lung alveoli, capillaries stratified: skin, mouth, vagina
cuboidal cube shaped, central nucleus glands, renal tubules
columnar tall, narrow, nucleus toward base tall, narrow, nucleus along cell simple: digestive tract pseudostratified: respiratory tract
transitional round, simple but appear stratified urinary bladder
Squamous Epithelia
Squamous epithelial cells are generally round, flat, and have a small, centrally located nucleus. The cell outline is slightly irregular, and cells fit together to form a covering or lining. When the cells are arranged in a single layer (simple epithelia), they facilitate diffusion in tissues, such as the areas of gas exchange in the lungs and the exchange of nutrients and waste at blood capillaries.
Figure \(1\) illustrates a layer of squamous cells with their membranes joined together to form an epithelium. Image Figure \(1\) illustrates squamous epithelial cells arranged in stratified layers, where protection is needed on the body from outside abrasion and damage. This is called a stratified squamous epithelium and occurs in the skin and in tissues lining the mouth and vagina.
Cuboidal Epithelia
Cuboidal epithelial cells, shown in Figure \(2\), are cube-shaped with a single, central nucleus. They are most commonly found in a single layer representing a simple epithelia in glandular tissues throughout the body where they prepare and secrete glandular material. They are also found in the walls of tubules and in the ducts of the kidney and liver.
Columnar Epithelia
Columnar epithelial cells are taller than they are wide: they resemble a stack of columns in an epithelial layer, and are most commonly found in a single-layer arrangement. The nuclei of columnar epithelial cells in the digestive tract appear to be lined up at the base of the cells, as illustrated in Figure \(3\). These cells absorb material from the lumen of the digestive tract and prepare it for entry into the body through the circulatory and lymphatic systems.
Columnar epithelial cells lining the respiratory tract appear to be stratified. However, each cell is attached to the base membrane of the tissue and, therefore, they are simple tissues. The nuclei are arranged at different levels in the layer of cells, making it appear as though there is more than one layer, as seen in Figure \(4\). This is called pseudostratified, columnar epithelia. This cellular covering has cilia at the apical, or free, surface of the cells. The cilia enhance the movement of mucous and trapped particles out of the respiratory tract, helping to protect the system from invasive microorganisms and harmful material that has been breathed into the body. Goblet cells are interspersed in some tissues (such as the lining of the trachea). The goblet cells contain mucous that traps irritants, which in the case of the trachea keep these irritants from getting into the lungs.
Transitional Epithelia
Transitional or uroepithelial cells appear only in the urinary system, primarily in the bladder and ureter. These cells are arranged in a stratified layer, but they have the capability of appearing to pile up on top of each other in a relaxed, empty bladder, as illustrated in Figure \(5\). As the urinary bladder fills, the epithelial layer unfolds and expands to hold the volume of urine introduced into it. As the bladder fills, it expands and the lining becomes thinner. In other words, the tissue transitions from thick to thin.
Exercise
Which of the following statements about types of epithelial cells is false?
1. Simple columnar epithelial cells line the tissue of the lung.
2. Simple cuboidal epithelial cells are involved in the filtering of blood in the kidney.
3. Pseudostratisfied columnar epithilia occur in a single layer, but the arrangement of nuclei makes it appear that more than one layer is present.
4. Transitional epithelia change in thickness depending on how full the bladder is.
Answer
A
Connective Tissues
Connective tissues are made up of a matrix consisting of living cells and a non-living substance, called the ground substance. The ground substance is made of an organic substance (usually a protein) and an inorganic substance (usually a mineral or water). The principal cell of connective tissues is the fibroblast. This cell makes the fibers found in nearly all of the connective tissues. Fibroblasts are motile, able to carry out mitosis, and can synthesize whichever connective tissue is needed. Macrophages, lymphocytes, and, occasionally, leukocytes can be found in some of the tissues. Some tissues have specialized cells that are not found in the others. The matrix in connective tissues gives the tissue its density. When a connective tissue has a high concentration of cells or fibers, it has proportionally a less dense matrix.
The organic portion or protein fibers found in connective tissues are either collagen, elastic, or reticular fibers. Collagen fibers provide strength to the tissue, preventing it from being torn or separated from the surrounding tissues. Elastic fibers are made of the protein elastin; this fiber can stretch to one and one half of its length and return to its original size and shape. Elastic fibers provide flexibility to the tissues. Reticular fibers are the third type of protein fiber found in connective tissues. This fiber consists of thin strands of collagen that form a network of fibers to support the tissue and other organs to which it is connected. The various types of connective tissues, the types of cells and fibers they are made of, and sample locations of the tissues is summarized in the table.
Table \(2\): Connective Tissues
Tissue Cells Fibers Location
loose/areolar fibroblasts, macrophages, some lymphocytes, some neutrophils few: collagen, elastic, reticular around blood vessels; anchors epithelia
dense, fibrous connective tissue fibroblasts, macrophages, mostly collagen irregular: skin regular: tendons, ligaments
cartilage chondrocytes, chondroblasts hyaline: few collagen fibrocartilage: large amount of collagen shark skeleton, fetal bones, human ears, intervertebral discs
bone osteoblasts, osteocytes, osteoclasts some: collagen, elastic vertebrate skeletons
adipose adipocytes few adipose (fat)
blood red blood cells, white blood cells none blood
Loose/Areolar Connective Tissue
Loose connective tissue, also called areolar connective tissue, has a sampling of all of the components of a connective tissue. As illustrated in Figure \(6\), loose connective tissue has some fibroblasts; macrophages are present as well. Collagen fibers are relatively wide and stain a light pink, while elastic fibers are thin and stain dark blue to black. The space between the formed elements of the tissue is filled with the matrix. The material in the connective tissue gives it a loose consistency similar to a cotton ball that has been pulled apart. Loose connective tissue is found around every blood vessel and helps to keep the vessel in place. The tissue is also found around and between most body organs. In summary, areolar tissue is tough, yet flexible, and comprises membranes.
Fibrous Connective Tissue
Fibrous connective tissues contain large amounts of collagen fibers and few cells or matrix material. The fibers can be arranged irregularly or regularly with the strands lined up in parallel. Irregularly arranged fibrous connective tissues are found in areas of the body where stress occurs from all directions, such as the dermis of the skin. Regular fibrous connective tissue, shown in Figure \(7\), is found in tendons (which connect muscles to bones) and ligaments (which connect bones to bones).
Cartilage
Cartilage is a connective tissue with a large amount of the matrix and variable amounts of fibers. The cells, called chondrocytes, make the matrix and fibers of the tissue. Chondrocytes are found in spaces within the tissue called lacunae.
A cartilage with few collagen and elastic fibers is hyaline cartilage, illustrated in Figure \(8\). The lacunae are randomly scattered throughout the tissue and the matrix takes on a milky or scrubbed appearance with routine histological stains. Sharks have cartilaginous skeletons, as does nearly the entire human skeleton during a specific pre-birth developmental stage. A remnant of this cartilage persists in the outer portion of the human nose. Hyaline cartilage is also found at the ends of long bones, reducing friction and cushioning the articulations of these bones.
Elastic cartilage has a large amount of elastic fibers, giving it tremendous flexibility. The ears of most vertebrate animals contain this cartilage as do portions of the larynx, or voice box. Fibrocartilage contains a large amount of collagen fibers, giving the tissue tremendous strength. Fibrocartilage comprises the intervertebral discs in vertebrate animals. Hyaline cartilage found in movable joints such as the knee and shoulder becomes damaged as a result of age or trauma. Damaged hyaline cartilage is replaced by fibrocartilage and results in the joints becoming “stiff.”
Bone
Bone, or osseous tissue, is a connective tissue that has a large amount of two different types of matrix material. The organic matrix is similar to the matrix material found in other connective tissues, including some amount of collagen and elastic fibers. This gives strength and flexibility to the tissue. The inorganic matrix consists of mineral salts—mostly calcium salts—that give the tissue hardness. Without adequate organic material in the matrix, the tissue breaks; without adequate inorganic material in the matrix, the tissue bends.
There are three types of cells in bone: osteoblasts, osteocytes, and osteoclasts. Osteoblasts are active in making bone for growth and remodeling. Osteoblasts deposit bone material into the matrix and, after the matrix surrounds them, they continue to live, but in a reduced metabolic state as osteocytes. Osteocytes are found in lacunae of the bone. Osteoclasts are active in breaking down bone for bone remodeling, and they provide access to calcium stored in tissues. Osteoclasts are usually found on the surface of the tissue.
Bone can be divided into two types: compact and spongy. Compact bone is found in the shaft (or diaphysis) of a long bone and the surface of the flat bones, while spongy bone is found in the end (or epiphysis) of a long bone. Compact bone is organized into subunits called osteons, as illustrated in Figure \(9\). A blood vessel and a nerve are found in the center of the structure within the Haversian canal, with radiating circles of lacunae around it known as lamellae. The wavy lines seen between the lacunae are microchannels called canaliculi; they connect the lacunae to aid diffusion between the cells. Spongy bone is made of tiny plates called trabeculae these plates serve as struts to give the spongy bone strength. Over time, these plates can break causing the bone to become less resilient. Bone tissue forms the internal skeleton of vertebrate animals, providing structure to the animal and points of attachment for tendons.
Adipose Tissue
Adipose tissue, or fat tissue, is considered a connective tissue even though it does not have fibroblasts or a real matrix and only has a few fibers. Adipose tissue is made up of cells called adipocytes that collect and store fat in the form of triglycerides, for energy metabolism. Adipose tissues additionally serve as insulation to help maintain body temperatures, allowing animals to be endothermic, and they function as cushioning against damage to body organs. Under a microscope, adipose tissue cells appear empty due to the extraction of fat during the processing of the material for viewing, as seen in Figure \(10\). The thin lines in the image are the cell membranes, and the nuclei are the small, black dots at the edges of the cells.
Blood
Blood is considered a connective tissue because it has a matrix, as shown in Figure \(11\). The living cell types are red blood cells (RBC), also called erythrocytes, and white blood cells (WBC), also called leukocytes. The fluid portion of whole blood, its matrix, is commonly called plasma.
The cell found in greatest abundance in blood is the erythrocyte. Erythrocytes are counted in millions in a blood sample: the average number of red blood cells in primates is 4.7 to 5.5 million cells per microliter. Erythrocytes are consistently the same size in a species, but vary in size between species. For example, the average diameter of a primate red blood cell is 7.5 µl, a dog is close at 7.0 µl, but a cat’s RBC diameter is 5.9 µl. Sheep erythrocytes are even smaller at 4.6 µl. Mammalian erythrocytes lose their nuclei and mitochondria when they are released from the bone marrow where they are made. Fish, amphibian, and avian red blood cells maintain their nuclei and mitochondria throughout the cell’s life. The principal job of an erythrocyte is to carry and deliver oxygen to the tissues.
Leukocytes are the predominant white blood cells found in the peripheral blood. Leukocytes are counted in the thousands in the blood with measurements expressed as ranges: primate counts range from 4,800 to 10,800 cells per µl, dogs from 5,600 to 19,200 cells per µl, cats from 8,000 to 25,000 cells per µl, cattle from 4,000 to 12,000 cells per µl, and pigs from 11,000 to 22,000 cells per µl.
Lymphocytes function primarily in the immune response to foreign antigens or material. Different types of lymphocytes make antibodies tailored to the foreign antigens and control the production of those antibodies. Neutrophils are phagocytic cells and they participate in one of the early lines of defense against microbial invaders, aiding in the removal of bacteria that has entered the body. Another leukocyte that is found in the peripheral blood is the monocyte. Monocytes give rise to phagocytic macrophages that clean up dead and damaged cells in the body, whether they are foreign or from the host animal. Two additional leukocytes in the blood are eosinophils and basophils—both help to facilitate the inflammatory response.
The slightly granular material among the cells is a cytoplasmic fragment of a cell in the bone marrow. This is called a platelet or thrombocyte. Platelets participate in the stages leading up to coagulation of the blood to stop bleeding through damaged blood vessels. Blood has a number of functions, but primarily it transports material through the body to bring nutrients to cells and remove waste material from them.
Muscle Tissues
There are three types of muscle in animal bodies: smooth, skeletal, and cardiac. They differ by the presence or absence of striations or bands, the number and location of nuclei, whether they are voluntarily or involuntarily controlled, and their location within the body. The table summarizes these differences.
Table \(3\): Types of Muscles
Type of Muscle Striations Nuclei Control Location
smooth no single, in center involuntary visceral organs
skeletal yes many, at periphery voluntary skeletal muscles
cardiac yes single, in center involuntary heart
Smooth Muscle
Smooth muscle does not have striations in its cells. It has a single, centrally located nucleus, as shown in Figure \(12\). Constriction of smooth muscle occurs under involuntary, autonomic nervous control and in response to local conditions in the tissues. Smooth muscle tissue is also called non-striated as it lacks the banded appearance of skeletal and cardiac muscle. The walls of blood vessels, the tubes of the digestive system, and the tubes of the reproductive systems are composed of mostly smooth muscle.
Skeletal Muscle
Skeletal muscle has striations across its cells caused by the arrangement of the contractile proteins actin and myosin. These muscle cells are relatively long and have multiple nuclei along the edge of the cell. Skeletal muscle is under voluntary, somatic nervous system control and is found in the muscles that move bones. Figure \(12\)illustrates the histology of skeletal muscle.
Cardiac Muscle
Cardiac muscle, shown in Figure \(12\), is found only in the heart. Like skeletal muscle, it has cross striations in its cells, but cardiac muscle has a single, centrally located nucleus. Cardiac muscle is not under voluntary control but can be influenced by the autonomic nervous system to speed up or slow down. An added feature to cardiac muscle cells is a line than extends along the end of the cell as it abuts the next cardiac cell in the row. This line is called an intercalated disc: it assists in passing electrical impulse efficiently from one cell to the next and maintains the strong connection between neighboring cardiac cells.
Nervous Tissues
Nervous tissues are made of cells specialized to receive and transmit electrical impulses from specific areas of the body and to send them to specific locations in the body. The main cell of the nervous system is the neuron, illustrated in Figure \(13\). The large structure with a central nucleus is the cell body of the neuron. Projections from the cell body are either dendrites specialized in receiving input or a single axon specialized in transmitting impulses. Some glial cells are also shown. Astrocytes regulate the chemical environment of the nerve cell, and oligodendrocytes insulate the axon so the electrical nerve impulse is transferred more efficiently. Other glial cells that are not shown support the nutritional and waste requirements of the neuron. Some of the glial cells are phagocytic and remove debris or damaged cells from the tissue. A nerve consists of neurons and glial cells.
Link to Learning
Click through the interactive review to learn more about epithelial tissues.
Career Connections: Pathologist
A pathologist is a medical doctor or veterinarian who has specialized in the laboratory detection of disease in animals, including humans. These professionals complete medical school education and follow it with an extensive post-graduate residency at a medical center. A pathologist may oversee clinical laboratories for the evaluation of body tissue and blood samples for the detection of disease or infection. They examine tissue specimens through a microscope to identify cancers and other diseases. Some pathologists perform autopsies to determine the cause of death and the progression of disease.
Summary
The basic building blocks of complex animals are four primary tissues. These are combined to form organs, which have a specific, specialized function within the body, such as the skin or kidney. Organs are organized together to perform common functions in the form of systems. The four primary tissues are epithelia, connective tissues, muscle tissues, and nervous tissues.
Glossary
canaliculus
microchannel that connects the lacunae and aids diffusion between cells
cartilage
type of connective tissue with a large amount of ground substance matrix, cells called chondrocytes, and some amount of fibers
chondrocyte
cell found in cartilage
columnar epithelia
epithelia made of cells taller than they are wide, specialized in absorption
connective tissue
type of tissue made of cells, ground substance matrix, and fibers
cuboidal epithelia
epithelia made of cube-shaped cells, specialized in glandular functions
epithelial tissue
tissue that either lines or covers organs or other tissues
fibrous connective tissue
type of connective tissue with a high concentration of fibers
lacuna
space in cartilage and bone that contains living cells
loose (areolar) connective tissue
type of connective tissue with small amounts of cells, matrix, and fibers; found around blood vessels
matrix
component of connective tissue made of both living and non-living (ground substances) cells
osteon
subunit of compact bone
pseudostratified
layer of epithelia that appears multilayered, but is a simple covering
simple epithelia
single layer of epithelial cells
squamous epithelia
type of epithelia made of flat cells, specialized in aiding diffusion or preventing abrasion
stratified epithelia
multiple layers of epithelial cells
trabecula
tiny plate that makes up spongy bone and gives it strength
transitional epithelia
epithelia that can transition for appearing multilayered to simple; also called uroepithelial | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/33%3A_The_Animal_Body_-_Basic_Form_and_Function/33.2%3A_Animal_Primary_Tissues.txt |
Skills to Develop
• Define homeostasis
• Describe the factors affecting homeostasis
• Discuss positive and negative feedback mechanisms used in homeostasis
• Describe thermoregulation of endothermic and ectothermic animals
Animal organs and organ systems constantly adjust to internal and external changes through a process called homeostasis (“steady state”). These changes might be in the level of glucose or calcium in blood or in external temperatures. Homeostasis means to maintain dynamic equilibrium in the body. It is dynamic because it is constantly adjusting to the changes that the body’s systems encounter. It is equilibrium because body functions are kept within specific ranges. Even an animal that is apparently inactive is maintaining this homeostatic equilibrium.
Homeostatic Process
The goal of homeostasis is the maintenance of equilibrium around a point or value called a set point. While there are normal fluctuations from the set point, the body’s systems will usually attempt to go back to this point. A change in the internal or external environment is called a stimulus and is detected by a receptor; the response of the system is to adjust the deviation parameter toward the set point. For instance, if the body becomes too warm, adjustments are made to cool the animal. If the blood’s glucose rises after a meal, adjustments are made to lower the blood glucose level by getting the nutrient into tissues that need it or to store it for later use.
Control of Homeostasis
When a change occurs in an animal’s environment, an adjustment must be made. The receptor senses the change in the environment, then sends a signal to the control center (in most cases, the brain) which in turn generates a response that is signaled to an effector. The effector is a muscle (that contracts or relaxes) or a gland that secretes. Homeostatsis is maintained by negative feedback loops. Positive feedback loops actually push the organism further out of homeostasis, but may be necessary for life to occur. Homeostasis is controlled by the nervous and endocrine system of mammals.
Negative Feedback Mechanisms
Any homeostatic process that changes the direction of the stimulus is a negative feedback loop. It may either increase or decrease the stimulus, but the stimulus is not allowed to continue as it did before the receptor sensed it. In other words, if a level is too high, the body does something to bring it down, and conversely, if a level is too low, the body does something to make it go up. Hence the term negative feedback. An example is animal maintenance of blood glucose levels. When an animal has eaten, blood glucose levels rise. This is sensed by the nervous system. Specialized cells in the pancreas sense this, and the hormone insulin is released by the endocrine system. Insulin causes blood glucose levels to decrease, as would be expected in a negative feedback system, as illustrated in Figure \(1\). However, if an animal has not eaten and blood glucose levels decrease, this is sensed in another group of cells in the pancreas, and the hormone glucagon is released causing glucose levels to increase. This is still a negative feedback loop, but not in the direction expected by the use of the term “negative.” Another example of an increase as a result of the feedback loop is the control of blood calcium. If calcium levels decrease, specialized cells in the parathyroid gland sense this and release parathyroid hormone (PTH), causing an increased absorption of calcium through the intestines and kidneys and, possibly, the breakdown of bone in order to liberate calcium. The effects of PTH are to raise blood levels of the element. Negative feedback loops are the predominant mechanism used in homeostasis.
Positive Feedback Loop
A positive feedback loop maintains the direction of the stimulus, possibly accelerating it. Few examples of positive feedback loops exist in animal bodies, but one is found in the cascade of chemical reactions that result in blood clotting, or coagulation. As one clotting factor is activated, it activates the next factor in sequence until a fibrin clot is achieved. The direction is maintained, not changed, so this is positive feedback. Another example of positive feedback is uterine contractions during childbirth, as illustrated in Figure \(2\). The hormone oxytocin, made by the endocrine system, stimulates the contraction of the uterus. This produces pain sensed by the nervous system. Instead of lowering the oxytocin and causing the pain to subside, more oxytocin is produced until the contractions are powerful enough to produce childbirth.
Exercise
State whether each of the following processes is regulated by a positive feedback loop or a negative feedback loop.
1. A person feels satiated after eating a large meal.
2. The blood has plenty of red blood cells. As a result, erythropoietin, a hormone that stimulates the production of new red blood cells, is no longer released from the kidney.
Answer
Both processes are the result of negative feedback loops. Negative feedback loops, which tend to keep a system at equilibrium, are more common than positive feedback loops.
Set Point
It is possible to adjust a system’s set point. When this happens, the feedback loop works to maintain the new setting. An example of this is blood pressure: over time, the normal or set point for blood pressure can increase as a result of continued increases in blood pressure. The body no longer recognizes the elevation as abnormal and no attempt is made to return to the lower set point. The result is the maintenance of an elevated blood pressure that can have harmful effects on the body. Medication can lower blood pressure and lower the set point in the system to a more healthy level. This is called a process of alteration of the set point in a feedback loop.
Changes can be made in a group of body organ systems in order to maintain a set point in another system. This is called acclimatization. This occurs, for instance, when an animal migrates to a higher altitude than it is accustomed to. In order to adjust to the lower oxygen levels at the new altitude, the body increases the number of red blood cells circulating in the blood to ensure adequate oxygen delivery to the tissues. Another example of acclimatization is animals that have seasonal changes in their coats: a heavier coat in the winter ensures adequate heat retention, and a light coat in summer assists in keeping body temperature from rising to harmful levels.
Link to Learning
Feedback mechanisms can be understood in terms of driving a race car along a track: watch a short video lesson on positive and negative feedback loops.
Homeostasis: Thermoregulation
Body temperature affects body activities. Generally, as body temperature rises, enzyme activity rises as well. For every ten degree centigrade rise in temperature, enzyme activity doubles, up to a point. Body proteins, including enzymes, begin to denature and lose their function with high heat (around 50oC for mammals). Enzyme activity will decrease by half for every ten degree centigrade drop in temperature, to the point of freezing, with a few exceptions. Some fish can withstand freezing solid and return to normal with thawing.
Link to Learning
Watch this Discovery Channel video on thermoregulation to see illustrations of this process in a variety of animals.
Endotherms and Ectotherms
Animals can be divided into two groups: some maintain a constant body temperature in the face of differing environmental temperatures, while others have a body temperature that is the same as their environment and thus varies with the environment. Animals that do not control their body temperature are ectotherms. This group has been called cold-blooded, but the term may not apply to an animal in the desert with a very warm body temperature. In contrast to ectotherms, which rely on external temperatures to set their body temperatures, poikilotherms are animals with constantly varying internal temperatures. An animal that maintains a constant body temperature in the face of environmental changes is called a homeotherm. Endotherms are animals that rely on internal sources for body temperature but which can exhibit extremes in temperature. These animals are able to maintain a level of activity at cooler temperature, which an ectotherm cannot due to differing enzyme levels of activity.
Heat can be exchanged between an animal and its environment through four mechanisms: radiation, evaporation, convection, and conduction (Figure \(3\)). Radiation is the emission of electromagnetic “heat” waves. Heat comes from the sun in this manner and radiates from dry skin the same way. Heat can be removed with liquid from a surface during evaporation. This occurs when a mammal sweats. Convection currents of air remove heat from the surface of dry skin as the air passes over it. Heat will be conducted from one surface to another during direct contact with the surfaces, such as an animal resting on a warm rock.
Heat Conservation and Dissipation
Animals conserve or dissipate heat in a variety of ways. In certain climates, endothermic animals have some form of insulation, such as fur, fat, feathers, or some combination thereof. Animals with thick fur or feathers create an insulating layer of air between their skin and internal organs. Polar bears and seals live and swim in a subfreezing environment and yet maintain a constant, warm, body temperature. The arctic fox, for example, uses its fluffy tail as extra insulation when it curls up to sleep in cold weather. Mammals have a residual effect from shivering and increased muscle activity: arrector pili muscles cause “goose bumps,” causing small hairs to stand up when the individual is cold; this has the intended effect of increasing body temperature. Mammals use layers of fat to achieve the same end. Loss of significant amounts of body fat will compromise an individual’s ability to conserve heat.
Endotherms use their circulatory systems to help maintain body temperature. Vasodilation brings more blood and heat to the body surface, facilitating radiation and evaporative heat loss, which helps to cool the body. Vasoconstriction reduces blood flow in peripheral blood vessels, forcing blood toward the core and the vital organs found there, and conserving heat. Some animals have adaptions to their circulatory system that enable them to transfer heat from arteries to veins, warming blood returning to the heart. This is called a countercurrent heat exchange; it prevents the cold venous blood from cooling the heart and other internal organs. This adaption can be shut down in some animals to prevent overheating the internal organs. The countercurrent adaption is found in many animals, including dolphins, sharks, bony fish, bees, and hummingbirds. In contrast, similar adaptations can help cool endotherms when needed, such as dolphin flukes and elephant ears.
Some ectothermic animals use changes in their behavior to help regulate body temperature. For example, a desert ectothermic animal may simply seek cooler areas during the hottest part of the day in the desert to keep from getting too warm. The same animals may climb onto rocks to capture heat during a cold desert night. Some animals seek water to aid evaporation in cooling them, as seen with reptiles. Other ectotherms use group activity such as the activity of bees to warm a hive to survive winter.
Many animals, especially mammals, use metabolic waste heat as a heat source. When muscles are contracted, most of the energy from the ATP used in muscle actions is wasted energy that translates into heat. Severe cold elicits a shivering reflex that generates heat for the body. Many species also have a type of adipose tissue called brown fat that specializes in generating heat.
Neural Control of Thermoregulation
The nervous system is important to thermoregulation, as illustrated in Figure \(4\). The processes of homeostasis and temperature control are centered in the hypothalamus of the advanced animal brain.
Exercise
When bacteria are destroyed by leuckocytes, pyrogens are released into the blood. Pyrogens reset the body’s thermostat to a higher temperature, resulting in fever. How might pyrogens cause the body temperature to rise?
Answer
Pyrogens increase body temperature by causing the blood vessels to constrict, inducing shivering, and stopping sweat glands from secreting fluid.
The hypothalamus maintains the set point for body temperature through reflexes that cause vasodilation and sweating when the body is too warm, or vasoconstriction and shivering when the body is too cold. It responds to chemicals from the body. When a bacterium is destroyed by phagocytic leukocytes, chemicals called endogenous pyrogens are released into the blood. These pyrogens circulate to the hypothalamus and reset the thermostat. This allows the body’s temperature to increase in what is commonly called a fever. An increase in body temperature causes iron to be conserved, which reduces a nutrient needed by bacteria. An increase in body heat also increases the activity of the animal’s enzymes and protective cells while inhibiting the enzymes and activity of the invading microorganisms. Finally, heat itself may also kill the pathogen. A fever that was once thought to be a complication of an infection is now understood to be a normal defense mechanism.
Summary
Homeostasis is a dynamic equilibrium that is maintained in body tissues and organs. It is dynamic because it is constantly adjusting to the changes that the systems encounter. It is in equilibrium because body functions are kept within a normal range, with some fluctuations around a set point for the processes.
Glossary
acclimatization
alteration in a body system in response to environmental change
alteration
change of the set point in a homeostatic system
homeostasis
dynamic equilibrium maintaining appropriate body functions
negative feedback loop
feedback to a control mechanism that increases or decreases a stimulus instead of maintaining it
positive feedback loop
feedback to a control mechanism that continues the direction of a stimulus
set point
midpoint or target point in homeostasis
thermoregulation
regulation of body temperature | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/33%3A_The_Animal_Body_-_Basic_Form_and_Function/33.3%3A_Homeostasis.txt |
33.1: Animal Form and Function
Animals vary in form and function. From a sponge to a worm to a goat, an organism has a distinct body plan that limits its size and shape. Animals’ bodies are also designed to interact with their environments, whether in the deep sea, a rainforest canopy, or the desert. Therefore, a large amount of information about the structure of an organism's body (anatomy) and the function of its cells, tissues and organs (physiology) can be learned by studying that organism's environment.
Review Questions
Which type of animal maintains a constant internal body temperature?
1. endotherm
2. ectotherm
3. coelomate
4. mesoderm
Answer
A
The symmetry found in animals that move swiftly is ________.
1. radial
2. bilateral
3. sequential
4. interrupted
Answer
B
What term describes the condition of a desert mouse that lowers its metabolic rate and “sleeps” during the hot day?
1. turgid
2. hibernation
3. estivation
4. normal sleep pattern
Answer
C
A plane that divides an animal into equal right and left portions is ________.
1. diagonal
2. midsagittal
3. coronal
4. transverse
Answer
B
A plane that divides an animal into dorsal and ventral portions is ________.
1. sagittal
2. midsagittal
3. coronal
4. transverse
Answer
D
The pleural cavity is a part of which cavity?
1. dorsal cavity
2. thoracic cavity
3. abdominal cavity
4. pericardial cavity
Answer
B
Free Response
How does diffusion limit the size of an organism? How is this counteracted?
Answer
Diffusion is effective over a very short distance. If a cell exceeds this distance in its size, the center of the cell cannot get adequate nutrients nor can it expel enough waste to survive. To compensate for this, cells can loosely adhere to each other in a liquid medium, or develop into multi-celled organisms that use circulatory and respiratory systems to deliver nutrients and remove wastes.
What is the relationship between BMR and body size? Why?
Answer
Basal Metabolic Rate is an expression of the metabolic processes that occur to maintain an individual’s functioning and body temperature. Smaller bodied animals have a relatively large surface area compared to a much larger animal. The large animal’s large surface area leads to increased heat loss that the animal must compensate for, resulting in a higher BMR. A small animal, having less relative surface area, does not lose as much heat and has a correspondingly lower BMR.
33.2: Animal Primary Tissues
The tissues of multicellular, complex animals are four primary types: epithelial, connective, muscle, and nervous. Recall that tissues are groups of similar cells group of similar cells carrying out related functions. These tissues combine to form organs—like the skin or kidney—that have specific, specialized functions within the body. Organs are organized into organ systems to perform functions.
Review Questions
Which type of epithelial cell is best adapted to aid diffusion?
1. squamous
2. cuboidal
3. columnar
4. transitional
Answer
C
Which type of epithelial cell is found in glands?
1. squamous
2. cuboidal
3. columnar
4. transitional
Answer
B
Which type of epithelial cell is found in the urinary bladder?
1. squamous
2. cuboidal
3. columnar
4. transitional
Answer
D
Which type of connective tissue has the most fibers?
1. loose connective tissue
2. fibrous connective tissue
3. cartilage
4. bone
Answer
B
Which type of connective tissue has a mineralized different matrix?
1. loose connective tissue
2. fibrous connective tissue
3. cartilage
4. bone
Answer
D
The cell found in bone that breaks it down is called an ________.
1. osteoblast
2. osteocyte
3. osteoclast
4. osteon
Answer
C
The cell found in bone that makes the bone is called an ________.
1. osteoblast
2. osteocyte
3. osteoclast
4. osteon
Answer
A
Plasma is the ________.
1. fibers in blood
2. matrix of blood
3. cell that phagocytizes bacteria
4. cell fragment found in the tissue
Answer
B
The type of muscle cell under voluntary control is the ________.
1. smooth muscle
2. skeletal muscle
3. cardiac muscle
4. visceral muscle
Answer
B
The part of a neuron that contains the nucleus is the
1. cell body
2. dendrite
3. axon
4. glial
Answer
B
Free Response
How can squamous epithelia both facilitate diffusion and prevent damage from abrasion?
Answer
Squamous epithelia can be either simple or stratified. As a single layer of cells, it presents a very thin epithelia that minimally inhibits diffusion. As a stratified epithelia, the surface cells can be sloughed off and the cells in deeper layers protect the underlying tissues from damage.
What are the similarities between cartilage and bone?
Answer
Both contain cells other than the traditional fibroblast. Both have cells that lodge in spaces within the tissue called lacunae. Both collagen and elastic fibers are found in bone and cartilage. Both tissues participate in vertebrate skeletal development and formation.
33.3: Homeostasis
Animal organs and organ systems constantly adjust to internal and external changes through a process called homeostasis (“steady state”). These changes might be in the level of glucose or calcium in blood or in external temperatures. Homeostasis means to maintain dynamic equilibrium in the body. It is dynamic because it is constantly adjusting to the changes that the body’s systems encounter. It is equilibrium because body functions are kept within specific ranges.
Review Questions
When faced with a sudden drop in environmental temperature, an endothermic animal will:
1. experience a drop in its body temperature
2. wait to see if it goes lower
3. increase muscle activity to generate heat
4. add fur or fat to increase insulation
Answer
C
Which is an example of negative feedback?
1. lowering of blood glucose after a meal
2. blood clotting after an injury
3. lactation during nursing
4. uterine contractions during labor
Answer
A
Which method of heat exchange occurs during direct contact between the source and animal?
1. radiation
2. evaporation
3. convection
4. conduction
Answer
D
The body’s thermostat is located in the ________.
1. homeostatic receptor
2. hypothalamus
3. medulla
4. vasodilation center
Answer
B
Free Response
Why are negative feedback loops used to control body homeostasis?
Answer
An adjustment to a change in the internal or external environment requires a change in the direction of the stimulus. A negative feedback loop accomplishes this, while a positive feedback loop would continue the stimulus and result in harm to the animal.
Why is a fever a “good thing” during a bacterial infection?
Answer
Mammalian enzymes increase activity to the point of denaturation, increasing the chemical activity of the cells involved. Bacterial enzymes have a specific temperature for their most efficient activity and are inhibited at either higher or lower temperatures. Fever results in an increase in the destruction of the invading bacteria by increasing the effectiveness of body defenses and an inhibiting bacterial metabolism.
How is a condition such as diabetes a good example of the failure of a set point in humans?
Answer
Diabetes is often associated with a lack in production of insulin. Without insulin, blood glucose levels go up after a meal, but never go back down to normal levels. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/33%3A_The_Animal_Body_-_Basic_Form_and_Function/33.E%3A_The_Animal_Body_-_Basic_Form_and_Function_%28Exercises%29.txt |
All living organisms need nutrients to survive. While plants can obtain the molecules required for cellular function through the process of photosynthesis, most animals obtain their nutrients by the consumption of other organisms. At the cellular level, the biological molecules necessary for animal function are amino acids, lipid molecules, nucleotides, and simple sugars. However, the food consumed consists of protein, fat, and complex carbohydrates. Animals must convert these macromolecules into the simple molecules required for maintaining cellular functions, such as assembling new molecules, cells, and tissues. The conversion of the food consumed to the nutrients required is a multi-step process involving digestion and absorption. During digestion, food particles are broken down to smaller components, and later, they are absorbed by the body.
• 34.0: Prelude to Animal Nutrition and the Digestive System
One of the challenges in human nutrition is maintaining a balance between food intake, storage, and energy expenditure. Imbalances can have serious health consequences. For example, eating too much food while not expending much energy leads to obesity, which in turn will increase the risk of developing illnesses such as type-2 diabetes and cardiovascular disease. The recent rise in obesity and related diseases makes understanding diet and nutrition important in maintaining good health.
• 34.1: 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.
• 34.2: Nutrition and Energy Production
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.
• 34.3: Digestive System Processes
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.
• 34.4: Digestive System Regulation
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.
• 34.E: Animal Nutrition and the Digestive System (Exercises)
Thumbnail: Intestine. (Image by JimCoote from Pixabay).
34: Animal Nutrition and the Digestive System
All living organisms need nutrients to survive. While plants can obtain the molecules required for cellular function through the process of photosynthesis, most animals obtain their nutrients by the consumption of other organisms. At the cellular level, the biological molecules necessary for animal function are amino acids, lipid molecules, nucleotides, and simple sugars. However, the food consumed consists of protein, fat, and complex carbohydrates. Animals must convert these macromolecules into the simple molecules required for maintaining cellular functions, such as assembling new molecules, cells, and tissues. The conversion of the food consumed to the nutrients required is a multi-step process involving digestion and absorption. During digestion, food particles are broken down to smaller components, and later, they are absorbed by the body.
One of the challenges in human nutrition is maintaining a balance between food intake, storage, and energy expenditure. Imbalances can have serious health consequences. For example, eating too much food while not expending much energy leads to obesity, which in turn will increase the risk of developing illnesses such as type-2 diabetes and cardiovascular disease. The recent rise in obesity and related diseases makes understanding the role of diet and nutrition in maintaining good health all the more important. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/34%3A_Animal_Nutrition_and_the_Digestive_System/34.0%3A_Prelude_to_Animal_Nutrition_and_the_Digestive_System.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/34%3A_Animal_Nutrition_and_the_Digestive_System/34.1%3A_Digestive_Systems.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/34%3A_Animal_Nutrition_and_the_Digestive_System/34.2%3A_Nutrition_and_Energy_Production.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/34%3A_Animal_Nutrition_and_the_Digestive_System/34.3%3A_Digestive_System_Processes.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/34%3A_Animal_Nutrition_and_the_Digestive_System/34.4%3A_Digestive_System_Regulation.txt |
34.1: 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.
Review Questions
Which of the following is a pseudo-ruminant?
1. cow
2. pig
3. crow
4. horse
Answer
D
Which of the following statements is untrue?
1. Roughage takes a long time to digest.
2. Birds eat large quantities at one time so that they can fly long distances.
3. Cows do not have upper teeth.
4. In pseudo-ruminants, roughage is digested in the cecum.
Answer
B
The acidic nature of chyme is neutralized by ________.
1. potassium hydroxide
2. sodium hydroxide
3. bicarbonates
4. vinegar
Answer
C
The digestive juices from the liver are delivered to the ________.
1. stomach
2. liver
3. duodenum
4. colon
Answer
C
Free Response
How does the polygastric digestive system aid in digesting roughage?
Answer
Animals with a polygastric digestive system have a multi-chambered stomach. The four compartments of the stomach are called the rumen, reticulum, omasum, and abomasum. These chambers contain many microbes that break down the cellulose and ferment the ingested food. The abomasum is the “true” stomach and is the equivalent of a monogastric stomach chamber where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the microbial support necessary for ruminants to digest plant material.
How do birds digest their food in the absence of teeth?
Answer
Birds have a stomach chamber called a gizzard. Here, the food is stored, soaked, and ground into finer particles, often using pebbles. Once this process is complete, the digestive juices take over in the proventriculus and continue the digestive process.
What is the role of the accessory organs in digestion?
Answer
Accessory organs play an important role in producing and delivering digestive juices to the intestine during digestion and absorption. Specifically, the salivary glands, liver, pancreas, and gallbladder play important roles. Malfunction of any of these organs can lead to disease states.
Explain how the villi and microvilli aid in absorption.
Answer
The villi and microvilli are folds on the surface of the small intestine. These folds increase the surface area of the intestine and provide more area for the absorption of nutrients.
34.2: Nutrition and Energy Production
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.
Review Questions
Which of the following statements is not true?
1. Essential nutrients can be synthesized by the body.
2. Vitamins are required in small quantities for bodily function.
3. Some amino acids can be synthesized by the body, while others need to be obtained from diet.
4. Vitamins come in two categories: fat-soluble and water-soluble.
Answer
A
Which of the following is a water-soluble vitamin?
1. vitamin A
2. vitamin E
3. vitamin K
4. vitamin C
Answer
D
What is the primary fuel for the body?
1. carbohydrates
2. lipids
3. protein
4. glycogen
Answer
A
Excess glucose is stored as ________.
1. fat
2. glucagon
3. glycogen
4. it is not stored in the body
Answer
C
Free Response
What are essential nutrients?
Answer
Essential nutrients are those nutrients that must be obtained from the diet because they cannot be produced by the body. Vitamins and minerals are examples of essential nutrients.
What is the role of minerals in maintaining good health?
Answer
Minerals—such as potassium, sodium, and calcium—are required for the functioning of many cellular processes, including muscle contraction and nerve conduction. While minerals are required in trace amounts, not having minerals in the diet can be potentially harmful.
Discuss why obesity is a growing epidemic.
Answer
In the United States, obesity, particularly childhood obesity, is a growing concern. Some of the contributors to this situation include sedentary lifestyles and consuming more processed foods and less fruits and vegetables. As a result, even young children who are obese can face health concerns.
There are several nations where malnourishment is a common occurrence. What may be some of the health challenges posed by malnutrition?
Answer
Malnutrition, often in the form of not getting enough calories or not enough of the essential nutrients, can have severe consequences. Many malnourished children have vision and dental problems, and over the years may develop many serious health problems.
34.3: Digestive System Processes
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.
Review Questions
Where does the majority of protein digestion take place?
1. stomach
2. duodenum
3. mouth
4. jejunum
Answer
A
Lipases are enzymes that break down ________.
1. disaccharides
2. lipids
3. proteins
4. cellulose
Answer
B
Free Response
Explain why some dietary lipid is a necessary part of a balanced diet.
Answer
Lipids add flavor to food and promote a sense of satiety or fullness. Fatty foods are sources of high energy; one gram of lipid contains nine calories. Lipids are also required in the diet to aid the absorption of lipid-soluble vitamins and for the production of lipid-soluble hormones.
34.4: Digestive System Regulation
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.
Review Questions
Which hormone controls the release of bile from the gallbladder
1. pepsin
2. amylase
3. CCK
4. gastrin
Answer
C
Which hormone stops acid secretion in the stomach?
1. gastrin
2. somatostatin
3. gastric inhibitory peptide
4. CCK
Answer
B
Free Response
Describe how hormones regulate digestion.
Answer
Hormones control the different digestive enzymes that are secreted in the stomach and the intestine during the process of digestion and absorption. For example, the hormone gastrin stimulates stomach acid secretion in response to food intake. The hormone somatostatin stops the release of stomach acid.
Describe one or more scenarios where loss of hormonal regulation of digestion can lead to diseases.
Answer
There are many cases where loss of hormonal regulation can lead to illnesses. For example, the bilirubin produced by the breakdown of red blood cells is converted to bile by the liver. When there is malfunction of this process, there is excess bilirubin in the blood and bile levels are low. As a result, the body struggles with dealing with fatty food. This is why a patient suffering from jaundice is asked to eat a diet with almost zero fat. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/34%3A_Animal_Nutrition_and_the_Digestive_System/34.E%3A_Animal_Nutrition_and_the_Digestive_System_%28Exercises%29.txt |
A nervous system is an organism’s control center: it processes sensory information from outside (and inside) the body and controls all behaviors—from eating to sleeping to finding a mate.
• 35.0: Prelude to the Nervous System
When you’re reading this book, your nervous system is performing several functions simultaneously. The visual system is processing what is seen on the page; the motor system controls the turn of the pages (or click of the mouse); the prefrontal cortex maintains attention. Even fundamental functions, like breathing and regulation of body temperature, are controlled by the nervous system.
• 35.1: Neurons and Glial Cells
Nervous systems throughout the animal kingdom vary in structure and complexity. 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.
• 35.2: How Neurons Communicate
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.
• 35.3: The Central Nervous System
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.
• 35.4: The Peripheral Nervous System
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).
• 35.5: Nervous System Disorders
A nervous system that functions correctly is a fantastically complex, well-oiled machine—synapses fire appropriately, muscles move when needed, memories are formed and stored, and emotions are well regulated. Unfortunately, each year millions of people in the United States deal with some sort of nervous system disorder.
• 35.E: The Nervous System (Exercises)
Thumbnail: Human brain toy. (Photo by Robina Weermeijer on Unsplash)
35: The Nervous System
When you’re reading this book, your nervous system is performing several functions simultaneously. The visual system is processing what is seen on the page; the motor system controls the turn of the pages (or click of the mouse); the prefrontal cortex maintains attention. Even fundamental functions, like breathing and regulation of body temperature, are controlled by the nervous system. A nervous system is an organism’s control center: it processes sensory information from outside (and inside) the body and controls all behaviors—from eating to sleeping to finding a mate. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.0%3A_Prelude_to_the_Nervous_System.txt |
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 | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.1%3A_Neurons_and_Glial_Cells.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 | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.2%3A_How_Neurons_Communicate.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.3%3A_The_Central_Nervous_System.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.4%3A_The_Peripheral_Nervous_System.txt |
Skills to Develop
• Describe the symptoms, potential causes, and treatment of several examples of nervous system disorders
A nervous system that functions correctly is a fantastically complex, well-oiled machine—synapses fire appropriately, muscles move when needed, memories are formed and stored, and emotions are well regulated. Unfortunately, each year millions of people in the United States deal with some sort of nervous system disorder. While scientists have discovered potential causes of many of these diseases, and viable treatments for some, ongoing research seeks to find ways to better prevent and treat all of these disorders.
Neurodegenerative Disorders
Neurodegenerative disorders are illnesses characterized by a loss of nervous system functioning that are usually caused by neuronal death. These diseases generally worsen over time as more and more neurons die. The symptoms of a particular neurodegenerative disease are related to where in the nervous system the death of neurons occurs. Spinocerebellar ataxia, for example, leads to neuronal death in the cerebellum. The death of these neurons causes problems in balance and walking. Neurodegenerative disorders include Huntington’s disease, amyotrophic lateral sclerosis, Alzheimer’s disease and other types of dementia disorders, and Parkinson’s disease. Here, Alzheimer’s and Parkinson’s disease will be discussed in more depth.
Alzheimer’s Disease
Alzheimer’s disease is the most common cause of dementia in the elderly. In 2012, an estimated 5.4 million Americans suffered from Alzheimer’s disease, and payments for their care are estimated at \$200 billion. Roughly one in every eight people age 65 or older has the disease. Due to the aging of the baby-boomer generation, there are projected to be as many as 13 million Alzheimer’s patients in the United States in the year 2050.
Symptoms of Alzheimer’s disease include disruptive memory loss, confusion about time or place, difficulty planning or executing tasks, poor judgment, and personality changes. Problems smelling certain scents can also be indicative of Alzheimer’s disease and may serve as an early warning sign. Many of these symptoms are also common in people who are aging normally, so it is the severity and longevity of the symptoms that determine whether a person is suffering from Alzheimer’s.
Alzheimer’s disease was named for Alois Alzheimer, a German psychiatrist who published a report in 1911 about a woman who showed severe dementia symptoms. Along with his colleagues, he examined the woman’s brain following her death and reported the presence of abnormal clumps, which are now called amyloid plaques, along with tangled brain fibers called neurofibrillary tangles. Amyloid plaques, neurofibrillary tangles, and an overall shrinking of brain volume are commonly seen in the brains of Alzheimer’s patients. Loss of neurons in the hippocampus is especially severe in advanced Alzheimer’s patients. Figure \(1\) compares a normal brain to the brain of an Alzheimer’s patient. Many research groups are examining the causes of these hallmarks of the disease.
One form of the disease is usually caused by mutations in one of three known genes. This rare form of early onset Alzheimer’s disease affects fewer than five percent of patients with the disease and causes dementia beginning between the ages of 30 and 60. The more prevalent, late-onset form of the disease likely also has a genetic component. One particular gene, apolipoprotein E (APOE) has a variant (E4) that increases a carrier’s likelihood of getting the disease. Many other genes have been identified that might be involved in the pathology.
Link to Learning
Visit this website for video links discussing genetics and Alzheimer’s disease.
Unfortunately, there is no cure for Alzheimer’s disease. Current treatments focus on managing the symptoms of the disease. Because decrease in the activity of cholinergic neurons (neurons that use the neurotransmitter acetylcholine) is common in Alzheimer’s disease, several drugs used to treat the disease work by increasing acetylcholine neurotransmission, often by inhibiting the enzyme that breaks down acetylcholine in the synaptic cleft. Other clinical interventions focus on behavioral therapies like psychotherapy, sensory therapy, and cognitive exercises. Since Alzheimer’s disease appears to hijack the normal aging process, research into prevention is prevalent. Smoking, obesity, and cardiovascular problems may be risk factors for the disease, so treatments for those may also help to prevent Alzheimer’s disease. Some studies have shown that people who remain intellectually active by playing games, reading, playing musical instruments, and being socially active in later life have a reduced risk of developing the disease.
Parkinson’s Disease
Like Alzheimer’s disease, Parkinson’s disease is a neurodegenerative disease. It was first characterized by James Parkinson in 1817. Each year, 50,000-60,000 people in the United States are diagnosed with the disease. Parkinson’s disease causes the loss of dopamine neurons in the substantia nigra, a midbrain structure that regulates movement. Loss of these neurons causes many symptoms including tremor (shaking of fingers or a limb), slowed movement, speech changes, balance and posture problems, and rigid muscles. The combination of these symptoms often causes a characteristic slow hunched shuffling walk, illustrated in Figure \(2\). Patients with Parkinson’s disease can also exhibit psychological symptoms, such as dementia or emotional problems.
Although some patients have a form of the disease known to be caused by a single mutation, for most patients the exact causes of Parkinson’s disease remain unknown: the disease likely results from a combination of genetic and environmental factors (similar to Alzheimer’s disease). Post-mortem analysis of brains from Parkinson’s patients shows the presence of Lewy bodies—abnormal protein clumps—in dopaminergic neurons. The prevalence of these Lewy bodies often correlates with the severity of the disease.
There is no cure for Parkinson’s disease, and treatment is focused on easing symptoms. One of the most commonly prescribed drugs for Parkinson’s is L-DOPA, which is a chemical that is converted into dopamine by neurons in the brain. This conversion increases the overall level of dopamine neurotransmission and can help compensate for the loss of dopaminergic neurons in the substantia nigra. Other drugs work by inhibiting the enzyme that breaks down dopamine.
Neurodevelopmental Disorders
Neurodevelopmental disorders occur when the development of the nervous system is disturbed. There are several different classes of neurodevelopmental disorders. Some, like Down Syndrome, cause intellectual deficits. Others specifically affect communication, learning, or the motor system. Some disorders like autism spectrum disorder and attention deficit/hyperactivity disorder have complex symptoms.
Autism
Autism spectrum disorder (ASD) is a neurodevelopmental disorder. Its severity differs from person to person. Estimates for the prevalence of the disorder have changed rapidly in the past few decades. Current estimates suggest that one in 88 children will develop the disorder. ASD is four times more prevalent in males than females.
A characteristic symptom of ASD is impaired social skills. Children with autism may have difficulty making and maintaining eye contact and reading social cues. They also may have problems feeling empathy for others. Other symptoms of ASD include repetitive motor behaviors (such as rocking back and forth), preoccupation with specific subjects, strict adherence to certain rituals, and unusual language use. Up to 30 percent of patients with ASD develop epilepsy, and patients with some forms of the disorder (like Fragile X) also have intellectual disability. Because it is a spectrum disorder, other ASD patients are very functional and have good-to-excellent language skills. Many of these patients do not feel that they suffer from a disorder and instead think that their brains just process information differently.
Except for some well-characterized, clearly genetic forms of autism (like Fragile X and Rett’s Syndrome), the causes of ASD are largely unknown. Variants of several genes correlate with the presence of ASD, but for any given patient, many different mutations in different genes may be required for the disease to develop. At a general level, ASD is thought to be a disease of “incorrect” wiring. Accordingly, brains of some ASD patients lack the same level of synaptic pruning that occurs in non-affected people. In the 1990s, a research paper linked autism to a common vaccine given to children. This paper was retracted when it was discovered that the author falsified data, and follow-up studies showed no connection between vaccines and autism.
Treatment for autism usually combines behavioral therapies and interventions, along with medications to treat other disorders common to people with autism (depression, anxiety, obsessive compulsive disorder). Although early interventions can help mitigate the effects of the disease, there is currently no cure for ASD.
Attention Deficit Hyperactivity Disorder (ADHD)
Approximately three to five percent of children and adults are affected by attention deficit/hyperactivity disorder (ADHD). Like ASD, ADHD is more prevalent in males than females. Symptoms of the disorder include inattention (lack of focus), executive functioning difficulties, impulsivity, and hyperactivity beyond what is characteristic of the normal developmental stage. Some patients do not have the hyperactive component of symptoms and are diagnosed with a subtype of ADHD: attention deficit disorder (ADD). Many people with ADHD also show comorbitity, in that they develop secondary disorders in addition to ADHD. Examples include depression or obsessive compulsive disorder (OCD). Figure \(3\) provides some statistics concerning comorbidity with ADHD.
The cause of ADHD is unknown, although research points to a delay and dysfunction in the development of the prefrontal cortex and disturbances in neurotransmission. According to studies of twins, the disorder has a strong genetic component. There are several candidate genes that may contribute to the disorder, but no definitive links have been discovered. Environmental factors, including exposure to certain pesticides, may also contribute to the development of ADHD in some patients. Treatment for ADHD often involves behavioral therapies and the prescription of stimulant medications, which paradoxically cause a calming effect in these patients.
Career Connection: Neurologist
Neurologists are physicians who specialize in disorders of the nervous system. They diagnose and treat disorders such as epilepsy, stroke, dementia, nervous system injuries, Parkinson’s disease, sleep disorders, and multiple sclerosis. Neurologists are medical doctors who have attended college, medical school, and completed three to four years of neurology residency.
When examining a new patient, a neurologist takes a full medical history and performs a complete physical exam. The physical exam contains specific tasks that are used to determine what areas of the brain, spinal cord, or peripheral nervous system may be damaged. For example, to check whether the hypoglossal nerve is functioning correctly, the neurologist will ask the patient to move his or her tongue in different ways. If the patient does not have full control over tongue movements, then the hypoglossal nerve may be damaged or there may be a lesion in the brainstem where the cell bodies of these neurons reside (or there could be damage to the tongue muscle itself).
Neurologists have other tools besides a physical exam they can use to diagnose particular problems in the nervous system. If the patient has had a seizure, for example, the neurologist can use electroencephalography (EEG), which involves taping electrodes to the scalp to record brain activity, to try to determine which brain regions are involved in the seizure. In suspected stroke patients, a neurologist can use a computerized tomography (CT) scan, which is a type of X-ray, to look for bleeding in the brain or a possible brain tumor. To treat patients with neurological problems, neurologists can prescribe medications or refer the patient to a neurosurgeon for surgery.
Link to Learning
This website allows you to see the different tests a neurologist might use to see what regions of the nervous system may be damaged in a patient.
Mental Illnesses
Mental illnesses are nervous system disorders that result in problems with thinking, mood, or relating with other people. These disorders are severe enough to affect a person’s quality of life and often make it difficult for people to perform the routine tasks of daily living. Debilitating mental disorders plague approximately 12.5 million Americans (about 1 in 17 people) at an annual cost of more than \$300 billion. There are several types of mental disorders including schizophrenia, major depression, bipolar disorder, anxiety disorders and phobias, post-traumatic stress disorders, and obsessive-compulsive disorder (OCD), among others. The American Psychiatric Association publishes the Diagnostic and Statistical Manual of Mental Disorders (or DSM), which describes the symptoms required for a patient to be diagnosed with a particular mental disorder. Each newly released version of the DSM contains different symptoms and classifications as scientists learn more about these disorders, their causes, and how they relate to each other. A more detailed discussion of two mental illnesses—schizophrenia and major depression—is given below.
Schizophrenia
Schizophrenia is a serious and often debilitating mental illness affecting one percent of people in the United States. Symptoms of the disease include the inability to differentiate between reality and imagination, inappropriate and unregulated emotional responses, difficulty thinking, and problems with social situations. People with schizophrenia can suffer from hallucinations and hear voices; they may also suffer from delusions. Patients also have so-called “negative” symptoms like a flattened emotional state, loss of pleasure, and loss of basic drives. Many schizophrenic patients are diagnosed in their late adolescence or early 20s. The development of schizophrenia is thought to involve malfunctioning dopaminergic neurons and may also involve problems with glutamate signaling. Treatment for the disease usually requires antipsychotic medications that work by blocking dopamine receptors and decreasing dopamine neurotransmission in the brain. This decrease in dopamine can cause Parkinson’s disease-like symptoms in some patients. While some classes of antipsychotics can be quite effective at treating the disease, they are not a cure, and most patients must remain medicated for the rest of their lives.
Depression
Major depression affects approximately 6.7 percent of the adults in the United States each year and is one of the most common mental disorders. To be diagnosed with major depressive disorder, a person must have experienced a severely depressed mood lasting longer than two weeks along with other symptoms including a loss of enjoyment in activities that were previously enjoyed, changes in appetite and sleep schedules, difficulty concentrating, feelings of worthlessness, and suicidal thoughts. The exact causes of major depression are unknown and likely include both genetic and environmental risk factors. Some research supports the “classic monoamine hypothesis,” which suggests that depression is caused by a decrease in norepinephrine and serotonin neurotransmission. One argument against this hypothesis is the fact that some antidepressant medications cause an increase in norepinephrine and serotonin release within a few hours of beginning treatment—but clinical results of these medications are not seen until weeks later. This has led to alternative hypotheses: for example, dopamine may also be decreased in depressed patients, or it may actually be an increase in norepinephrine and serotonin that causes the disease, and antidepressants force a feedback loop that decreases this release. Treatments for depression include psychotherapy, electroconvulsive therapy, deep-brain stimulation, and prescription medications. There are several classes of antidepressant medications that work through different mechanisms. For example, monoamine oxidase inhibitors (MAO inhibitors) block the enzyme that degrades many neurotransmitters (including dopamine, serotonin, norepinephrine), resulting in increased neurotransmitter in the synaptic cleft. Selective serotonin reuptake inhibitors (SSRIs) block the reuptake of serotonin into the presynaptic neuron. This blockage results in an increase in serotonin in the synaptic cleft. Other types of drugs such as norepinephrine-dopamine reuptake inhibitors and norepinephrine-serotonin reuptake inhibitors are also used to treat depression.
Other Neurological Disorders
There are several other neurological disorders that cannot be easily placed in the above categories. These include chronic pain conditions, cancers of the nervous system, epilepsy disorders, and stroke. Epilepsy and stroke are discussed below.
Epilepsy
Estimates suggest that up to three percent of people in the United States will be diagnosed with epilepsy in their lifetime. While there are several different types of epilepsy, all are characterized by recurrent seizures. Epilepsy itself can be a symptom of a brain injury, disease, or other illness. For example, people who have intellectual disability or ASD can experience seizures, presumably because the developmental wiring malfunctions that caused their disorders also put them at risk for epilepsy. For many patients, however, the cause of their epilepsy is never identified and is likely to be a combination of genetic and environmental factors. Often, seizures can be controlled with anticonvulsant medications. However, for very severe cases, patients may undergo brain surgery to remove the brain area where seizures originate.
Stroke
A stroke results when blood fails to reach a portion of the brain for a long enough time to cause damage. Without the oxygen supplied by blood flow, neurons in this brain region die. This neuronal death can cause many different symptoms—depending on the brain area affected— including headache, muscle weakness or paralysis, speech disturbances, sensory problems, memory loss, and confusion. Stroke is often caused by blood clots and can also be caused by the bursting of a weak blood vessel. Strokes are extremely common and are the third most common cause of death in the United States. On average one person experiences a stroke every 40 seconds in the United States. Approximately 75 percent of strokes occur in people older than 65. Risk factors for stroke include high blood pressure, diabetes, high cholesterol, and a family history of stroke. Smoking doubles the risk of stroke. Because a stroke is a medical emergency, patients with symptoms of a stroke should immediately go to the emergency room, where they can receive drugs that will dissolve any clot that may have formed. These drugs will not work if the stroke was caused by a burst blood vessel or if the stroke occurred more than three hours before arriving at the hospital. Treatment following a stroke can include blood pressure medication (to prevent future strokes) and (sometimes intense) physical therapy.
Summary
Some general themes emerge from the sampling of nervous system disorders presented above. The causes for most disorders are not fully understood—at least not for all patients—and likely involve a combination of nature (genetic mutations that become risk factors) and nurture (emotional trauma, stress, hazardous chemical exposure). Because the causes have yet to be fully determined, treatment options are often lacking and only address symptoms.
Glossary
Alzheimer’s disease
neurodegenerative disorder characterized by problems with memory and thinking
attention deficit hyperactivity disorder (ADHD)
neurodevelopmental disorder characterized by difficulty maintaining attention and controlling impulses
autism spectrum disorder (ASD)
neurodevelopmental disorder characterized by impaired social interaction and communication abilities
epilepsy
neurological disorder characterized by recurrent seizures
major depression
mental illness characterized by prolonged periods of sadness
neurodegenerative disorder
nervous system disorder characterized by the progressive loss of neurological functioning, usually caused by neuron death
Parkinson’s disease
neurodegenerative disorder that affects the control of movement
schizophrenia
mental disorder characterized by the inability to accurately perceive reality; patients often have difficulty thinking clearly and can suffer from delusions | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.5%3A_Nervous_System_Disorders.txt |
35.1: Neurons and Glial Cells
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.
Review Questions
Neurons contain ________, which can receive signals from other neurons.
1. axons
2. mitochondria
3. dendrites
4. Golgi bodies
Answer
C
A(n) ________ neuron has one axon and one dendrite extending directly from the cell body.
1. unipolar
2. bipolar
3. multipolar
4. pseudounipolar
Answer
B
Glia that provide myelin for neurons in the brain are called ________.
1. Schwann cells
2. oligodendrocytes
3. microglia
4. astrocytes
Answer
B
Free Response
How are neurons similar to other cells? How are they unique?
Answer
Neurons contain organelles common to all cells, such as a nucleus and mitochondria. They are unique because they contain dendrites, which can receive signals from other neurons, and axons that can send these signals to other cells.
Multiple sclerosis causes demyelination of axons in the brain and spinal cord. Why is this problematic?
Answer
Myelin provides insulation for signals traveling along axons. Without myelin, signal transmission can slow down and degrade over time. This would slow down neuronal communication across the nervous system and affect all downstream functions.
35.2: How Neurons Communicate
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.
Review Questions
For a neuron to fire an action potential, its membrane must reach ________.
1. hyperpolarization
2. the threshold of excitation
3. the refractory period
4. inhibitory postsynaptic potential
Answer
B
After an action potential, the opening of additional voltage-gated ________ channels and the inactivation of sodium channels, cause the membrane to return to its resting membrane potential.
1. sodium
2. potassium
3. calcium
4. chloride
Answer
B
What is the term for protein channels that connect two neurons at an electrical synapse?
1. synaptic vesicles
2. voltage-gated ion channels
3. gap junction protein
4. sodium-potassium exchange pumps
Answer
C
Free Response
How does myelin aid propagation of an action potential along an axon? How do the nodes of Ranvier help this process?
Answer
Myelin prevents the leak of current from the axon. Nodes of Ranvier allow the action potential to be regenerated at specific points along the axon. They also save energy for the cell since voltage-gated ion channels and sodium-potassium transporters are not needed along myelinated portions of the axon.
What are the main steps in chemical neurotransmission?
Answer
An action potential travels along an axon until it depolarizes the membrane at an axon terminal. Depolarization of the membrane causes voltage-gated Ca2+ channels to open and Ca2+ to enter the cell. The intracellular calcium influx causes synaptic vesicles containing neurotransmitter to fuse with the presynaptic membrane. The neurotransmitter diffuses across the synaptic cleft and binds to receptors on the postsynaptic membrane. Depending on the specific neurotransmitter and postsynaptic receptor, this action can cause positive (excitatory postsynaptic potential) or negative (inhibitory postsynaptic potential) ions to enter the cell.
35.3: The Central Nervous System
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.
Review Questions
The ________ lobe contains the visual cortex.
1. frontal
2. parietal
3. temporal
4. occipital
Answer
D
The ________ connects the two cerebral hemispheres.
1. limbic system
2. corpus callosum
3. cerebellum
4. pituitary
Answer
B
Neurons in the ________ control motor reflexes.
1. thalamus
2. spinal cord
3. parietal lobe
4. hippocampus
Answer
B
Free Response
What methods can be used to determine the function of a particular brain region?
Answer
To determine the function of a specific brain area, scientists can look at patients who have damage in that brain area and see what symptoms they exhibit. Researchers can disable the brain structure temporarily using transcranial magnetic stimulation. They can disable or remove the area in an animal model. fMRI can be used to correlate specific functions with increased blood flow to brain regions.
What are the main functions of the spinal cord?
Answer
The spinal cord transmits sensory information from the body to the brain and motor commands from the brain to the body through its connections with peripheral nerves. It also controls motor reflexes.
35.4: The Peripheral Nervous System
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).
Review Questions
Activation of the sympathetic nervous system causes:
1. increased blood flow into the skin
2. a decreased heart rate
3. an increased heart rate
4. increased digestion
Answer
C
Where are parasympathetic preganglionic cell bodies located?
1. cerebellum
2. brainstem
3. dorsal root ganglia
4. skin
Answer
B
________ is released by motor nerve endings onto muscle.
1. Acetylcholine
2. Norepinephrine
3. Dopamine
4. Serotonin
Answer
A
Free Response
What are the main differences between the sympathetic and parasympathetic branches of the autonomic nervous system?
Answer
The sympathetic nervous system prepares the body for “fight or flight,” whereas the parasympathetic nervous system allows the body to “rest and digest.” Sympathetic neurons release norepinephrine onto target organs; parasympathetic neurons release acetylcholine. Sympathetic neuron cell bodies are located in sympathetic ganglia. Parasympathetic neuron cell bodies are located in the brainstem and sacral spinal cord. Activation of the sympathetic nervous system increases heart rate and blood pressure and decreases digestion and blood flow to the skin. Activation of the parasympathetic nervous system decreases heart rate and blood pressure and increases digestion and blood flow to the skin.
What are the main functions of the sensory-somatic nervous system?
Answer
The sensory-somatic nervous system transmits sensory information from the skin, muscles, and sensory organs to the CNS. It also sends motor commands from the CNS to the muscles, causing them to contract.
35.5: Nervous System Disorders
A nervous system that functions correctly is a fantastically complex, well-oiled machine—synapses fire appropriately, muscles move when needed, memories are formed and stored, and emotions are well regulated. Unfortunately, each year millions of people in the United States deal with some sort of nervous system disorder.
Review Questions
Parkinson’s disease is a caused by the degeneration of neurons that release ________.
1. serotonin
2. dopamine
3. glutamate
4. norepinephrine
Answer
B
________ medications are often used to treat patients with ADHD.
1. Tranquilizer
2. Antibiotic
3. Stimulant
4. Anti-seizure
Answer
C
Strokes are often caused by ________.
1. neurodegeneration
2. blood clots or burst blood vessels
3. seizures
4. viruses
Answer
B
Free Response
What are the main symptoms of Alzheimer’s disease?
Answer
Symptoms of Alzheimer’s disease include disruptive memory loss, confusion about time or place, difficulties planning or executing tasks, poor judgment, and personality changes.
What are possible treatments for patients with major depression?
Answer
Possible treatments for patients with major depression include psychotherapy and prescription medications. MAO inhibitor drugs inhibit the breakdown of certain neurotransmitters (including dopamine, serotonin, norepinephrine) in the synaptic cleft. SSRI medications inhibit the reuptake of serotonin into the presynaptic neuron. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/35%3A_The_Nervous_System/35.E%3A_The_Nervous_System_%28Exercises%29.txt |
In more advanced animals, the senses are constantly at work, making the animal aware of stimuli—such as light, or sound, or the presence of a chemical substance in the external environment—and monitoring information about the organism’s internal environment. All bilaterally symmetric animals have a sensory system, and the development of any species’ sensory system has been driven by natural selection; thus, sensory systems differ among species according to the demands of their environments.
• 36.0: Introduction
The shark, unlike most fish predators, is electrosensitive—that is, sensitive to electrical fields produced by other animals in its environment. While it is helpful to this underwater predator, electrosensitivity is a sense not found in most land animals.
• 36.1: Sensory Processes
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.
• 36.2: Somatosensation
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.
• 36.3: Taste and Smell
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.
• 36.4: Hearing and Vestibular Sensation
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.
• 36.5: 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.
• 36.E: Sensory Systems (Exercises)
Thumbnail: Owl eyes. (Image by Graham Hobster from Pixabay).
36: Sensory Systems
In more advanced animals, the senses are constantly at work, making the animal aware of stimuli—such as light, or sound, or the presence of a chemical substance in the external environment—and monitoring information about the organism’s internal environment. All bilaterally symmetric animals have a sensory system, and the development of any species’ sensory system has been driven by natural selection; thus, sensory systems differ among species according to the demands of their environments. The shark, unlike most fish predators, is electrosensitive—that is, sensitive to electrical fields produced by other animals in its environment. While it is helpful to this underwater predator, electrosensitivity is a sense not found in most land animals. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.0%3A_Introduction.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.1%3A_Sensory_Processes.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.2%3A_Somatosensation.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.3%3A_Taste_and_Smell.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.4%3A_Hearing_and_Vestibular_Sensation.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.5%3A_Vision.txt |
36.1: Sensory Processes
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.
Review Questions
Where does perception occur?
1. spinal cord
2. cerebral cortex
3. receptors
4. thalamus
Answer
B
If a person’s cold receptors no longer convert cold stimuli into sensory signals, that person has a problem with the process of ________.
1. reception
2. transmission
3. perception
4. transduction
Answer
D
After somatosensory transduction, the sensory signal travels through the brain as a(n) _____ signal.
1. electrical
2. pressure
3. optical
4. thermal
Answer
A
Free Response
If a person sustains damage to axons leading from sensory receptors to the central nervous system, which step or steps of sensory perception will be affected?
Answer
Transmission of sensory information from the receptor to the central nervous system will be impaired, and thus, perception of stimuli, which occurs in the brain, will be halted.
In what way does the overall magnitude of a stimulus affect the just-noticeable difference in the perception of that stimulus?
Answer
The just-noticeable difference is a fraction of the overall magnitude of the stimulus and seems to be a relatively fixed proportion (such as 10 percent) whether the stimulus is large (such as a very heavy object) or small (such as a very light object).
36.2: Somatosensation
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.
Review Questions
_____ are found only in _____ skin, and detect skin deflection.
1. Meissner’s corpuscles: hairy
2. Merkel’s disks: glabrous
3. hair receptors: hairy
4. Krause end bulbs: hairy
Answer
B
If you were to burn your epidermis, what receptor type would you most likely burn?
1. free nerve endings
2. Ruffini endings
3. Pacinian corpuscle
4. hair receptors
Answer
A
Free Response
What can be inferred about the relative sizes of the areas of cortex that process signals from skin not densely innervated with sensory receptors and skin that is densely innervated with sensory receptors?
Answer
The cortical areas serving skin that is densely innervated likely are larger than those serving skin that is less densely innervated.
36.3: Taste and Smell
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.
Review Questions
Which of the following has the fewest taste receptors?
1. fungiform papillae
2. circumvallate papillae
3. foliate papillae
4. filiform papillae
Answer
D
How many different taste molecules do taste cells each detect?
1. one
2. five
3. ten
4. It depends on the spot on the tongue
Answer
A
Salty foods activate the taste cells by _____.
1. exciting the taste cell directly
2. causing hydrogen ions to enter the cell
3. causing sodium channels to close
4. binding directly to the receptors
Answer
A
All sensory signals except _____ travel to the _____ in the brain before the cerebral cortex.
1. vision; thalamus
2. olfaction; thalamus
3. vision; cranial nerves
4. olfaction; cranial nerves
Answer
B
Free Response
From the perspective of the recipient of the signal, in what ways do pheromones differ from other odorants?
Answer
Pheromones may not be consciously perceived, and pheromones can have direct physiological and behavioral effects on their recipients.
What might be the effect on an animal of not being able to perceive taste?
Answer
The animal might not be able to recognize the differences in food sources and thus might not be able to discriminate between spoiled food and safe food or between foods that contain necessary nutrients, such as proteins, and foods that do not.
36.4: Hearing and Vestibular Sensation
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.
Review Questions
In sound, pitch is measured in _____, and volume is measured in _____.
1. nanometers (nm); decibels (dB)
2. decibels (dB); nanometers (nm)
3. decibels (dB); hertz (Hz)
4. hertz (Hz); decibels (dB)
Answer
D
Auditory hair cells are indirectly anchored to the _____.
1. basilar membrane
2. oval window
3. tectorial membrane
4. ossicles
Answer
A
Which of the following are found both in the auditory system and the vestibular system?
1. basilar membrane
2. hair cells
3. semicircular canals
4. ossicles
Answer
B
Free Response
How would a rise in altitude likely affect the speed of a sound transmitted through air? Why?
Answer
The sound would slow down, because it is transmitted through the particles (gas) and there are fewer particles (lower density) at higher altitudes.
How might being in a place with less gravity than Earth has (such as Earth’s moon) affect vestibular sensation, and why?
Answer
Because vestibular sensation relies on gravity’s effects on tiny crystals in the inner ear, a situation of reduced gravity would likely impair vestibular sensation.
36.5: 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.
Review Questions
Why do people over 55 often need reading glasses?
1. Their cornea no longer focuses correctly.
2. Their lens no longer focuses correctly.
3. Their eyeball has elongated with age, causing images to focus in front of their retina.
4. Their retina has thinned with age, making vision more difficult.
Answer
B
Why is it easier to see images at night using peripheral, rather than the central, vision?
1. Cones are denser in the periphery of the retina.
2. Bipolar cells are denser in the periphery of the retina.
3. Rods are denser in the periphery of the retina.
4. The optic nerve exits at the periphery of the retina.
Answer
C
A person catching a ball must coordinate her head and eyes. What part of the brain is helping to do this?
1. hypothalamus
2. pineal gland
3. thalamus
4. superior colliculus
Answer
D
Free Response
How could the pineal gland, the brain structure that plays a role in annual cycles, use visual information from the suprachiasmatic nucleus of the hypothalamus?
Answer
The pineal gland could use length-of-day information to determine the time of year, for example. Day length is shorter in the winter than it is in the summer. For many animals and plants, photoperiod cues them to reproduce at a certain time of year.
How is the relationship between photoreceptors and bipolar cells different from other sensory receptors and adjacent cells?
Answer
The photoreceptors tonically inhibit the bipolar cells, and stimulation of the receptors turns this inhibition off, activating the bipolar cells. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/36%3A_Sensory_Systems/36.E%3A_Sensory_Systems_%28Exercises%29.txt |
An animal’s endocrine system controls body processes through the production, secretion, and regulation of hormones, which serve as chemical “messengers” functioning in cellular and organ activity and, ultimately, maintaining the body’s homeostasis. The endocrine system plays a role in growth, metabolism, and sexual development. In humans, common endocrine system diseases include thyroid disease and diabetes mellitus. In organisms that undergo metamorphosis, the process is controlled by the endocrine system. The transformation from tadpole to frog, for example, is complex and nuanced to adapt to specific environments and ecological circumstances.
• 37.0: Introduction
An animal’s endocrine system controls body processes through the production, secretion, and regulation of hormones, which serve as chemical “messengers” functioning in cellular and organ activity and, ultimately, maintaining the body’s homeostasis. The endocrine system plays a role in growth, metabolism, and sexual development. In humans, common endocrine system diseases include thyroid disease and diabetes mellitus.
• 37.1: Types of Hormones
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.
• 37.2: How Hormones Work
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.
• 37.3: Regulation of Body Processes
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.
• 37.4: Regulation of Hormone Production
Hormone production and release are primarily controlled by negative feedback. In negative feedback systems, a stimulus elicits the release of a substance; once the substance reaches a certain level, it sends a signal that stops further release of the substance. In this way, the concentration of hormones in blood is maintained within a narrow range.
• 37.5: Endocrine Glands
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.
• 37.E: The Endocrine System (Exercises)
Thumbnail: Wood frog tadpole. (CC BY 2.0; Brian Gratwicke via Flickr).
37: The Endocrine System
An animal’s endocrine system controls body processes through the production, secretion, and regulation of hormones, which serve as chemical “messengers” functioning in cellular and organ activity and, ultimately, maintaining the body’s homeostasis. The endocrine system plays a role in growth, metabolism, and sexual development. In humans, common endocrine system diseases include thyroid disease and diabetes mellitus. In organisms that undergo metamorphosis, the process is controlled by the endocrine system. The transformation from tadpole to frog, for example, is complex and nuanced to adapt to specific environments and ecological circumstances. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.0%3A_Introduction.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 | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.1%3A_Types_of_Hormones.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.2%3A_How_Hormones_Work.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.3%3A_Regulation_of_Body_Processes.txt |
Skills to Develop
• Explain how hormone production is regulated
• Discuss the different stimuli that control hormone levels in the body
Hormone production and release are primarily controlled by negative feedback. In negative feedback systems, a stimulus elicits the release of a substance; once the substance reaches a certain level, it sends a signal that stops further release of the substance. In this way, the concentration of hormones in blood is maintained within a narrow range. For example, the anterior pituitary signals the thyroid to release thyroid hormones. Increasing levels of these hormones in the blood then give feedback to the hypothalamus and anterior pituitary to inhibit further signaling to the thyroid gland, as illustrated in Figure \(1\). There are three mechanisms by which endocrine glands are stimulated to synthesize and release hormones: humoral stimuli, hormonal stimuli, and neural stimuli.
Exercise
Hyperthyroidism is a condition in which the thyroid gland is overactive. Hypothyroidism is a condition in which the thyroid gland is underactive. Which of the conditions are the following two patients most likely to have?
• Patient A has symptoms including weight gain, cold sensitivity, low heart rate and fatigue.
• Patient B has symptoms including weight loss, profuse sweating, increased heart rate and difficulty sleeping.
Answer
Patient A has symptoms associated with decreased metabolism, and may be suffering from hypothyroidism. Patient B has symptoms associated with increased metabolism, and may be suffering from hyperthyroidism.
Humoral Stimuli
The term “humoral” is derived from the term “humor,” which refers to bodily fluids such as blood. A humoral stimulus refers to the control of hormone release in response to changes in extracellular fluids such as blood or the ion concentration in the blood. For example, a rise in blood glucose levels triggers the pancreatic release of insulin. Insulin causes blood glucose levels to drop, which signals the pancreas to stop producing insulin in a negative feedback loop.
Hormonal Stimuli
Hormonal stimuli refers to the release of a hormone in response to another hormone. A number of endocrine glands release hormones when stimulated by hormones released by other endocrine glands. For example, the hypothalamus produces hormones that stimulate the anterior portion of the pituitary gland. The anterior pituitary in turn releases hormones that regulate hormone production by other endocrine glands. The anterior pituitary releases the thyroid-stimulating hormone, which then stimulates the thyroid gland to produce the hormones T3 and T4. As blood concentrations of T3 and T4 rise, they inhibit both the pituitary and the hypothalamus in a negative feedback loop.
Neural Stimuli
In some cases, the nervous system directly stimulates endocrine glands to release hormones, which is referred to as neural stimuli. Recall that in a short-term stress response, the hormones epinephrine and norepinephrine are important for providing the bursts of energy required for the body to respond. Here, neuronal signaling from the sympathetic nervous system directly stimulates the adrenal medulla to release the hormones epinephrine and norepinephrine in response to stress.
Summary
Hormone levels are primarily controlled through negative feedback, in which rising levels of a hormone inhibit its further release. The three mechanisms of hormonal release are humoral stimuli, hormonal stimuli, and neural stimuli. Humoral stimuli refers to the control of hormonal release in response to changes in extracellular fluid levels or ion levels. Hormonal stimuli refers to the release of hormones in response to hormones released by other endocrine glands. Neural stimuli refers to the release of hormones in response to neural stimulation.
Glossary
hormonal stimuli
release of a hormone in response to another hormone
humoral stimuli
control of hormone release in response to changes in extracellular fluids such as blood or the ion concentration in the blood
neural stimuli
stimulation of endocrine glands by the nervous system | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.4%3A_Regulation_of_Hormone_Production.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.5%3A_Endocrine_Glands.txt |
37.1: Types of Hormones
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.
Review Questions
A newly discovered hormone contains four amino acids linked together. Under which chemical class would this hormone be classified?
1. lipid-derived hormone
2. amino acid-derived hormone
3. peptide hormone
4. glycoprotein
Answer
C
Which class of hormones can diffuse through plasma membranes?
1. lipid-derived hormones
2. amino acid-derived hormones
3. peptide hormones
4. glycoprotein hormones
Answer
A
Free Response
Although there are many different hormones in the human body, they can be divided into three classes based on their chemical structure. What are these classes and what is one factor that distinguishes them?
Answer
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 hormones. One of the key distinguishing features of the lipid-derived hormones is that they can diffuse across plasma membranes whereas the amino acid-derived and peptide hormones cannot.
Where is insulin stored, and why would it be released?
Answer
Secreted peptides such as 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.
37.2: How Hormones Work
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.
Review Questions
A new antagonist molecule has been discovered that binds to and blocks plasma membrane receptors. What effect will this antagonist have on testosterone, a steroid hormone?
1. It will block testosterone from binding to its receptor.
2. It will block testosterone from activating cAMP signaling.
3. It will increase testosterone-mediated signaling.
4. It will not affect testosterone-mediated signaling.
Answer
D
What effect will a cAMP inhibitor have on a peptide hormone-mediated signaling pathway?
1. It will prevent the hormone from binding its receptor.
2. It will prevent activation of a G-protein.
3. It will prevent activation of adenylate cyclase.
4. It will prevent activation of protein kinases.
Answer
D
Free Response
Name two important functions of hormone receptors.
Answer
The number of receptors that respond to a hormone can change, resulting in increased or decreased cell sensitivity. The number of receptors can increase in response to rising hormone levels, called up-regulation, making the cell more sensitive to the hormone and allowing for more cellular activity. The number of receptors can also decrease in response to rising hormone levels, called down-regulation, leading to reduced cellular activity.
How can hormones mediate changes?
Answer
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 receptors and modulating gene transcription, or indirectly by binding to cell surface receptors and stimulating signaling pathways.
37.3: Regulation of Body Processes
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.
Review Questions
Drinking alcoholic beverages causes an increase in urine output. This most likely occurs because alcohol:
1. inhibits ADH release
2. stimulates ADH release
3. inhibits TSH release
4. stimulates TSH release
Answer
A
FSH and LH release from the anterior pituitary is stimulated by ________.
1. TSH
2. GnRH
3. T3
4. PTH
Answer
B
What hormone is produced by beta cells of the pancreas?
1. T3
2. glucagon
3. insulin
4. T4
Answer
C
When blood calcium levels are low, PTH stimulates:
1. excretion of calcium from the kidneys
2. excretion of calcium from the intestines
3. osteoblasts
4. osteoclasts
Answer
D
Free Response
Name and describe a function of one hormone produced by the anterior pituitary and one hormone produced by the posterior pituitary.
Answer
In addition to producing FSH and LH, the anterior pituitary 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 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 vagina stimulates oxytocin release in childbirth. Contractions increase in intensity as blood levels of oxytocin rise until the birth is complete.
Describe one direct action of growth hormone (GH).
Answer
Hormonal regulation is required for the growth and replication of most cells in the body. Growth hormone (GH), produced by the anterior pituitary, accelerates the rate of protein synthesis, particularly in skeletal muscles and bones. Growth hormone has direct and indirect mechanisms of action. The direct actions of GH include: 1) stimulation of fat 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. 2) In the liver, GH stimulates glycogen breakdown, which 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.
37.4: Regulation of Hormone Production
Hormone production and release are primarily controlled by negative feedback. In negative feedback systems, a stimulus elicits the release of a substance; once the substance reaches a certain level, it sends a signal that stops further release of the substance. In this way, the concentration of hormones in blood is maintained within a narrow range.
Review Questions
A rise in blood glucose levels triggers release of insulin from the pancreas. This mechanism of hormone production is stimulated by:
1. humoral stimuli
2. hormonal stimuli
3. neural stimuli
4. negative stimuli
Answer
A
Which mechanism of hormonal stimulation would be affected if signaling and hormone release from the hypothalamus was blocked?
1. humoral and hormonal stimuli
2. hormonal and neural stimuli
3. neural and humoral stimuli
4. hormonal and negative stimuli
Answer
B
Free Response
How is hormone production and release primarily controlled?
Answer
Hormone production and release are primarily controlled by negative feedback. In negative feedback systems, a stimulus causes the release of a substance whose effects then inhibit further release. In this way, the concentration of hormones in blood is maintained within a narrow range. For example, the anterior pituitary signals the thyroid to release thyroid hormones. Increasing levels of these hormones in the blood then feed back to the hypothalamus and anterior pituitary to inhibit further signaling to the thyroid gland.
Compare and contrast hormonal and humoral stimuli.
Answer
The term humoral is derived from the term humor, which refers to bodily fluids such as blood. Humoral stimuli refer to the control of hormone release in response to changes in extracellular fluids such as blood or the ion concentration in the blood. For example, a rise in blood glucose levels triggers the pancreatic release of insulin. Insulin causes blood glucose levels to drop, which signals the pancreas to stop producing insulin in a negative feedback loop.
Hormonal stimuli refer to the release of a hormone in response to another hormone. A number of endocrine glands release hormones when stimulated by hormones released by other endocrine organs. For example, the hypothalamus produces hormones that stimulate the anterior pituitary. The anterior pituitary in turn releases hormones that regulate hormone production by other endocrine glands. For example, the anterior pituitary releases thyroid-stimulating hormone, which stimulates the thyroid gland to produce the hormones T3 and T4. As blood concentrations of T3 and T4 rise they inhibit both the pituitary and the hypothalamus in a negative feedback loop.
37.5: Endocrine Glands
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.
Review Questions
Which endocrine glands are associated with the kidneys?
1. thyroid glands
2. pituitary glands
3. adrenal glands
4. gonads
Answer
C
Which of the following hormones is not produced by the anterior pituitary?
1. oxytocin
2. growth hormone
3. prolactin
4. thyroid-stimulating hormone
Answer
A
Free Response
What does aldosterone regulate, and how is it stimulated?
Answer
The main mineralocorticoid is aldosterone, which regulates the concentration of ions in urine, sweat, 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 an increase in blood potassium levels.
The adrenal medulla contains two types of secretory cells, what are they and what are their functions?
Answer
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 75–80 percent of its secretions. Epinephrine and norepinephrine increase heart rate, breathing rate, cardiac muscle contractions, 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. These neural impulses originate from the hypothalamus in response to stress to prepare the body for the fight-or-flight response. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/37%3A_The_Endocrine_System/37.E%3A_The_Endocrine_System_%28Exercises%29.txt |
The muscular and skeletal systems provide support to the body and allow for a wide range of movement. The bones of the skeletal system protect the body’s internal organs and support the weight of the body. The muscles of the muscular system contract and pull on the bones, allowing for movements as diverse as standing, walking, running, and grasping items.
• 38.0: Prelude to the Musculoskeletal System
Injury or disease affecting the musculoskeletal system can be very debilitating. In humans, the most common musculoskeletal diseases worldwide are caused by malnutrition. Ailments that affect the joints are also widespread, such as arthritis, which can make movement difficult and—in advanced cases—completely impair mobility. In severe cases in which the joint has suffered extensive damage, joint replacement surgery may be needed.
• 38.1: 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.
• 38.2: 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.
• 38.3: Joints and 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.
• 38.4: Muscle Contraction and Locomotion
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.
• 38.E: The Musculoskeletal System (Exercises)
Thumbnail: Muscular anatomical torso. (Photo by Alan Calvert on Unsplash)
38: The Musculoskeletal System
The muscular and skeletal systems provide support to the body and allow for a wide range of movement. The bones of the skeletal system protect the body’s internal organs and support the weight of the body. The muscles of the muscular system contract and pull on the bones, allowing for movements as diverse as standing, walking, running, and grasping items.
Injury or disease affecting the musculoskeletal system can be very debilitating. In humans, the most common musculoskeletal diseases worldwide are caused by malnutrition. Ailments that affect the joints are also widespread, such as arthritis, which can make movement difficult and—in advanced cases—completely impair mobility. In severe cases in which the joint has suffered extensive damage, joint replacement surgery may be needed.
Progress in the science of prosthesis design has resulted in the development of artificial joints, with joint replacement surgery in the hips and knees being the most common. Replacement joints for shoulders, elbows, and fingers are also available. Even with this progress, there is still room for improvement in the design of prostheses. The state-of-the-art prostheses have limited durability and therefore wear out quickly, particularly in young or active individuals. Current research is focused on the use of new materials, such as carbon fiber, that may make prostheses more durable. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/38%3A_The_Musculoskeletal_System/38.0%3A_Prelude_to_the_Musculoskeletal_System.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/38%3A_The_Musculoskeletal_System/38.1%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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/38%3A_The_Musculoskeletal_System/38.2%3A_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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/38%3A_The_Musculoskeletal_System/38.3%3A_Joints_and_Skeletal_Movement.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/38%3A_The_Musculoskeletal_System/38.4%3A_Muscle_Contraction_and_Locomotion.txt |
38.1: Types of Skeletal Systems
Review Questions
The forearm consists of the:
1. radius and ulna
2. radius and humerus
3. ulna and humerus
4. humerus and carpus
Answer
A
The pectoral girdle consists of the:
1. clavicle and sternum
2. sternum and scapula
3. clavicle and scapula
4. clavicle and coccyx
Answer
C
All of the following are groups of vertebrae except ________, which is a curvature.
1. thoracic
2. cervical
3. lumbar
4. pelvic
Answer
D
Which of these is a facial bone?
1. frontal
2. occipital
3. lacrimal
4. temporal
Answer
C
Free Response
What are the major differences between the male pelvis and female pelvis that permit childbirth in females?
Answer
The female pelvis is tilted forward and is wider, lighter, and shallower than the male pelvis. It is also has a pubic angle that is broader than the male pelvis.
What are the major differences between the pelvic girdle and the pectoral girdle that allow the pelvic girdle to bear the weight of the body?
Answer
The pelvic girdle is securely attached to the body by strong ligaments, unlike the pectoral girdle, which is sparingly attached to the ribcage. The sockets of the pelvic girdle are deep, allowing the femur to be more stable than the pectoral girdle, which has shallow sockets for the scapula. Most tetrapods have 75 percent of their weight on the front legs because the head and neck are so heavy; the advantage of the shoulder joint is more degrees of freedom in movement.
38.2: Bone
Review Questions
The Haversian canal:
1. is arranged as rods or plates
2. contains the bone’s blood vessels and nerve fibers
3. is responsible for the lengthwise growth of long bones
4. synthesizes and secretes matrix
Answer
B
The epiphyseal plate:
1. is arranged as rods or plates
2. contains the bone’s blood vessels and nerve fibers
3. is responsible for the lengthwise growth of long bones
4. synthesizes and secretes bone matrix
Answer
C
The cells responsible for bone resorption are ________.
1. osteoclasts
2. osteoblasts
3. fibroblasts
4. osteocytes
Answer
A
Compact bone is composed of ________.
1. trabeculae
2. compacted collagen
3. osteons
4. calcium phosphate only
Answer
C
Free Response
What are the major differences between spongy bone and compact bone?
Answer
Compact bone tissue forms the hard external layer of all bones and consists of osteons. Compact bone tissue is prominent in areas of bone at which stresses are applied in only a few directions. Spongy bone tissue forms the inner layer of all bones and consists of trabeculae. Spongy bone is prominent in areas of bones that are not heavily stressed or at which stresses arrive from many directions.
What are the roles of osteoblasts, osteocytes, and osteoclasts?
Answer
Osteocytes function in the exchange of nutrients and wastes with the blood. They also maintain normal bone structure by recycling the mineral salts in the bony matrix. Osteoclasts remove bone tissue by releasing lysosomal enzymes and acids that dissolve the bony matrix. Osteoblasts are bone cells that are responsible for bone formation.
38.3: Joints and Skeletal Movement
Review Questions
Synchondroses and symphyses are:
1. synovial joints
2. cartilaginous joints
3. fibrous joints
4. condyloid joints
Answer
B
The movement of bone away from the midline of the body is called ________.
1. circumduction
2. extension
3. adduction
4. abduction
Answer
D
Which of the following is not a characteristic of the synovial fluid?
1. lubrication
2. shock absorption
3. regulation of water balance in the joint
4. protection of articular cartilage
Answer
C
The elbow is an example of which type of joint?
1. hinge
2. pivot
3. saddle
4. gliding
Answer
A
Free Response
What movements occur at the hip joint and knees as you bend down to touch your toes?
Answer
The hip joint is flexed and the knees are extended.
What movement(s) occur(s) at the scapulae when you shrug your shoulders?
Answer
Elevation is the movement of a bone upward, such as when the shoulders are shrugged, lifting the scapulae. Depression is the downward movement of a bone, such as after the shoulders are shrugged and the scapulae return to their normal position from an elevated position.
38.4: Muscle Contraction and Locomotion
Review Questions
In relaxed muscle, the myosin-binding site on actin is blocked by ________.
1. titin
2. troponin
3. myoglobin
4. tropomyosin
Answer
D
The cell membrane of a muscle fiber is called a ________.
1. myofibril
2. sarcolemma
3. sarcoplasm
4. myofilament
Answer
B
The muscle relaxes if no new nerve signal arrives. However the neurotransmitter from the previous stimulation is still present in the synapse. The activity of ________ helps to remove this neurotransmitter.
1. myosin
2. action potential
3. tropomyosin
4. acetylcholinesterase
Answer
D
The ability of a muscle to generate tension immediately after stimulation is dependent on:
1. myosin interaction with the M line
2. overlap of myosin and actin
3. actin attachments to the Z line
4. none of the above
Answer
D
Free Response
How would muscle contractions be affected if ATP was completely depleted in a muscle fiber?
Answer
Because ATP is required for myosin to release from actin, muscles would remain rigidly contracted until more ATP was available for the myosin cross-bridge release. This is why dead vertebrates undergo rigor mortis.
What factors contribute to the amount of tension produced in an individual muscle fiber?
Answer
The cross-sectional area, the length of the muscle fiber at rest, and the frequency of neural stimulation.
What effect will low blood calcium have on neurons? What effect will low blood calcium have on skeletal muscles?
Answer
Neurons will not be able to release neurotransmitter without calcium. Skeletal muscles have calcium stored and don’t need any from the outside. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/38%3A_The_Musculoskeletal_System/New_Page.txt |
Breathing is an involuntary event. How often a breath is taken and how much air is inhaled or exhaled are tightly regulated by the respiratory center in the brain. Humans, when they aren’t exerting themselves, breathe approximately 15 times per minute on average. Canines have a respiratory rate of about 15–30 breaths per minute. With every inhalation, air fills the lungs, and with every exhalation, air rushes back out. That air is doing more than just inflating and deflating the lungs in the chest cavity. The air contains oxygen that crosses the lung tissue, enters the bloodstream, and travels to organs and tissues.
• 39.0: Prelude to the Respiratory System
The air contains oxygen that crosses the lung tissue, enters the bloodstream, and travels to organs and tissues. Oxygen enters the cells where it is used for metabolic reactions that produce ATP, a high-energy compound. At the same time, these reactions release carbon dioxide as a by-product. Carbon dioxide is toxic and must be eliminated. Carbon dioxide exits the cells, enters the bloodstream, travels back to the lungs, and is expired out of the body during exhalation.
• 39.1: Systems of Gas Exchange
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.
• 39.2: Gas Exchange across Respiratory Surfaces
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.
• 39.3: Breathing
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.
• 39.4: Transport of Gases in Human Bodily Fluids
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.
• 39.E: The Respiratory System (Exercises)
39: The Respiratory System
Breathing is an involuntary event. How often a breath is taken and how much air is inhaled or exhaled are tightly regulated by the respiratory center in the brain. Humans, when they aren’t exerting themselves, breathe approximately 15 times per minute on average. Canines have a respiratory rate of about 15–30 breaths per minute. With every inhalation, air fills the lungs, and with every exhalation, air rushes back out. That air is doing more than just inflating and deflating the lungs in the chest cavity. The air contains oxygen that crosses the lung tissue, enters the bloodstream, and travels to organs and tissues. Oxygen (O2) enters the cells where it is used for metabolic reactions that produce ATP, a high-energy compound. At the same time, these reactions release carbon dioxide (CO2) as a by-product. CO2 is toxic and must be eliminated. Carbon dioxide exits the cells, enters the bloodstream, travels back to the lungs, and is expired out of the body during exhalation. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/39%3A_The_Respiratory_System/39.0%3A_Prelude_to_the_Respiratory_System.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/39%3A_The_Respiratory_System/39.1%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 | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/39%3A_The_Respiratory_System/39.2%3A_Gas_Exchange_across_Respiratory_Surfaces.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/39%3A_The_Respiratory_System/39.3%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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/39%3A_The_Respiratory_System/39.4%3A_Transport_of_Gases_in_Human_Bodily_Fluids.txt |
39.1: Systems of Gas Exchange
Review Questions
The respiratory system ________.
1. provides body tissues with oxygen
2. provides body tissues with oxygen and carbon dioxide
3. establishes how many breaths are taken per minute
4. provides the body with carbon dioxide
Answer
A
Air is warmed and humidified in the nasal passages. This helps to ________.
1. ward off infection
2. decrease sensitivity during breathing
3. prevent damage to the lungs
4. all of the above
Answer
C
Which is the order of airflow during inhalation?
1. nasal cavity, trachea, larynx, bronchi, bronchioles, alveoli
2. nasal cavity, larynx, trachea, bronchi, bronchioles, alveoli
3. nasal cavity, larynx, trachea, bronchioles, bronchi, alveoli
4. nasal cavity, trachea, larynx, bronchi, bronchioles, alveoli
Answer
B
Free Response
Describe the function of these terms and describe where they are located: main bronchus, trachea, alveoli, and acinus.
Answer
The main bronchus is the conduit in the lung that funnels air to the airways where gas exchange occurs. The main bronchus attaches the lungs to the very end of the trachea where it bifurcates. The trachea is the cartilaginous structure that extends from the pharynx to the primary bronchi. It serves to funnel air to the lungs. The alveoli are the sites of gas exchange; they are located at the terminal regions of the lung and are attached to the respiratory bronchioles. The acinus is the structure in the lung where gas exchange occurs.
How does the structure of alveoli maximize gas exchange?
Answer
The sac-like structure of the alveoli increases their surface area. In addition, the alveoli are made of thin-walled parenchymal cells. These features allow gases to easily diffuse across the cells.
39.2: Gas Exchange across Respiratory Surfaces
Review Questions
The inspiratory reserve volume measures the ________.
1. amount of air remaining in the lung after a maximal exhalation
2. amount of air that the lung holds
3. amount of air the can be further exhaled after a normal breath
4. amount of air that can be further inhaled after a normal breath
Answer
D
Of the following, which does not explain why the partial pressure of oxygen is lower in the lung than in the external air?
1. Air in the lung is humidified; therefore, water vapor pressure alters the pressure.
2. Carbon dioxide mixes with oxygen.
3. Oxygen is moved into the blood and is headed to the tissues.
4. Lungs exert a pressure on the air to reduce the oxygen pressure.
Answer
D
The total lung capacity is calculated using which of the following formulas?
1. residual volume + tidal volume + inspiratory reserve volume
2. residual volume + expiratory reserve volume + inspiratory reserve volume
3. expiratory reserve volume + tidal volume + inspiratory reserve volume
4. residual volume + expiratory reserve volume + tidal volume + inspiratory reserve volume
Answer
D
Free Response
What does FEV1/FVC measure? What factors may affect FEV1/FVC?
Answer
FEV1/FVC measures the forced expiratory volume in one second in relation to the total forced vital capacity (the total amount of air that is exhaled from the lung from a maximal inhalation). This ratio changes with alterations in lung function that arise from diseases such as fibrosis, asthma, and COPD.
What is the reason for having residual volume in the lung?
Answer
If all the air in the lung were exhaled, then opening the alveoli for the next inspiration would be very difficult. This is because the tissues would stick together.
How can a decrease in the percent of oxygen in the air affect the movement of oxygen in the body?
Answer
Oxygen moves from the lung to the bloodstream to the tissues according to the pressure gradient. This is measured as the partial pressure of oxygen. If the amount of oxygen drops in the inspired air, there would be reduced partial pressure. This would decrease the driving force that moves the oxygen into the blood and into the tissues. $\text{P}_{\text{O}_2}$ is also reduced at high elevations: $\text{P}_{\text{O}_2}$ at high elevations is lower than at sea level because the total atmospheric pressure is less than atmospheric pressure at sea level.
If a patient has increased resistance in his or her lungs, how can this detected by a doctor? What does this mean?
Answer
A doctor can detect a restrictive disease using spirometry. By detecting the rate at which air can be expelled from the lung, a diagnosis of fibrosis or another restrictive disease can be made.
39.3: Breathing
Review Questions
How would paralysis of the diaphragm alter inspiration?
1. It would prevent contraction of the intercostal muscles.
2. It would prevent inhalation because the intrapleural pressure would not change.
3. It would decrease the intrapleural pressure and allow more air to enter the lungs.
4. It would slow expiration because the lung would not relax.
Answer
B
Restrictive airway diseases ________.
1. increase the compliance of the lung
2. decrease the compliance of the lung
3. increase the lung volume
4. decrease the work of breathing
Answer
B
Alveolar ventilation remains constant when ________.
1. the respiratory rate is increased while the volume of air per breath is decreased
2. the respiratory rate and the volume of air per breath are increased
3. the respiratory rate is decreased while increasing the volume per breath
4. both a and c
Answer
D
Free Response
How would increased airway resistance affect intrapleural pressure during inhalation?
Answer
Increased airway resistance increases the volume and pressure in the lung; therefore, the intrapleural pressure would be less negative and breathing would be more difficult.
Explain how a puncture to the thoracic cavity (from a knife wound, for instance) could alter the ability to inhale.
Answer
A puncture to the thoracic cavity would equalize the pressure inside the thoracic cavity to the outside environment. For the lung to function properly, the intrapleural pressure must be negative. This is caused by the contraction of the diaphragm pulling the lungs down and drawing air into the lungs.
When someone is standing, gravity stretches the bottom of the lung down toward the floor to a greater extent than the top of the lung. What implication could this have on the flow of air in the lungs? Where does gas exchange occur in the lungs?
Answer
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.
39.4: Transport of Gases in Human Bodily Fluids
Review Questions
Which of the following will NOT facilitate the transfer of oxygen to tissues?
1. decreased body temperature
2. decreased pH of the blood
3. increased carbon dioxide
4. increased exercise
Answer
A
The majority of carbon dioxide in the blood is transported by ________.
1. binding to hemoglobin
2. dissolution in the blood
3. conversion to bicarbonate
4. binding to plasma proteins
Answer
C
The majority of oxygen in the blood is transported by ________.
1. dissolution in the blood
2. being carried as bicarbonate ions
3. binding to blood plasma
4. binding to hemoglobin
Answer
D
Free Response
What would happen if no carbonic anhydrase were present in red blood cells?
Answer
Without carbonic anhydrase, carbon dioxide would not be hydrolyzed into carbonic acid or bicarbonate. Therefore, very little carbon dioxide (only 15 percent) would be transported in the blood away from the tissues.
How does the administration of 100 percent oxygen save a patient from carbon monoxide poisoning? Why wouldn’t giving carbon dioxide work?
Answer
Carbon monoxide has a higher affinity for hemoglobin than oxygen. This means that carbon monoxide will preferentially bind to hemoglobin over oxygen. Administration of 100 percent oxygen is an effective therapy because at that concentration, oxygen will displace the carbon monoxide from the hemoglobin. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/39%3A_The_Respiratory_System/39.E%3A_The_Respiratory_System_%28Exercises%29.txt |
Most animals are complex multicellular organisms that require a mechanism for transporting nutrients throughout their bodies and removing waste products. The circulatory system has evolved over time from simple diffusion through cells in the early evolution of animals to a complex network of blood vessels that reach all parts of the human body. This extensive network supplies the cells, tissues, and organs with oxygen and nutrients, and removes carbon dioxide and waste, which are byproducts of respiration.
• 40.0: Prelude to the Circulatory System
Gas exchange is one essential function of the circulatory system. A circulatory system is not needed in organisms with no specialized respiratory organs because oxygen and carbon dioxide diffuse directly between their body tissues and the external environment. However, in organisms that possess lungs and gills, oxygen must be transported from these specialized respiratory organs to the body tissues via a circulatory system.
• 40.1: Overview of the 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.
• 40.2: Components of the 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.
• 40.3: Mammalian Heart and Blood Vessels
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.
• 40.4: Blood Flow and Blood Pressure Regulation
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.
• 40.E: The Circulatory System (Exercises)
Thumbnail: The human heart. (CC-BY 4.0 / modified from original; OpenStax).
40: The Circulatory System
Most animals are complex multicellular organisms that require a mechanism for transporting nutrients throughout their bodies and removing waste products. The circulatory system has evolved over time from simple diffusion through cells in the early evolution of animals to a complex network of blood vessels that reach all parts of the human body. This extensive network supplies the cells, tissues, and organs with oxygen and nutrients, and removes carbon dioxide and waste, which are byproducts of respiration.
At the core of the human circulatory system is the heart. The size of a clenched fist, the human heart is protected beneath the rib cage. Made of specialized and unique cardiac muscle, it pumps blood throughout the body and to the heart itself. Heart contractions are driven by intrinsic electrical impulses that the brain and endocrine hormones help to regulate. Understanding the heart’s basic anatomy and function is important to understanding the body’s circulatory and respiratory systems.
Gas exchange is one essential function of the circulatory system. A circulatory system is not needed in organisms with no specialized respiratory organs because oxygen and carbon dioxide diffuse directly between their body tissues and the external environment. However, in organisms that possess lungs and gills, oxygen must be transported from these specialized respiratory organs to the body tissues via a circulatory system. Therefore, circulatory systems have had to evolve to accommodate the great diversity of body sizes and body types present among animals. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/40%3A_The_Circulatory_System/40.0%3A_Prelude_to_the_Circulatory_System.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/40%3A_The_Circulatory_System/40.1%3A_Overview_of_the_Circulatory_System.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/40%3A_The_Circulatory_System/40.2%3A_Components_of_the_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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/40%3A_The_Circulatory_System/40.3%3A_Mammalian_Heart_and_Blood_Vessels.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/40%3A_The_Circulatory_System/40.4%3A_Blood_Flow_and_Blood_Pressure_Regulation.txt |
40.1: Overview of the 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.
Review Questions
Why are open circulatory systems advantageous to some animals?
1. They use less metabolic energy.
2. They help the animal move faster.
3. They do not need a heart.
4. They help large insects develop.
Answer
A
Some animals use diffusion instead of a circulatory system. Examples include:
1. birds and jellyfish
2. flatworms and arthropods
3. mollusks and jellyfish
4. None of the above
Answer
D
Blood flow that is directed through the lungs and back to the heart is called ________.
1. unidirectional circulation
2. gill circulation
3. pulmonary circulation
4. pulmocutaneous circulation
Answer
C
Free Response
Describe a closed circulatory system.
Answer
A closed circulatory system is a closed-loop system, in which blood is not free in a cavity. Blood is separate from the bodily interstitial fluid and contained within blood vessels. In this type of system, blood circulates unidirectionally from the heart around the systemic circulatory route, and then returns to the heart.
Describe systemic circulation.
Answer
Systemic circulation flows through the systems of the body. The blood flows away from the heart to the brain, liver, kidneys, stomach, and other organs, the limbs, and the muscles of the body; it then returns to the heart.
40.2: Components of the 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.
Review Questions
White blood cells:
1. can be classified as granulocytes or agranulocytes
2. defend the body against bacteria and viruses
3. are also called leucocytes
4. All of the above
Answer
D
Platelet plug formation occurs at which point?
1. when large megakaryocytes break up into thousands of smaller fragments
2. when platelets are dispersed through the blood stream
3. when platelets are attracted to a site of blood vessel damage
4. none of the above
Answer
C
In humans, the plasma comprises what percentage of the blood?
1. 45 percent
2. 55 percent
3. 25 percent
4. 90 percent
Answer
B
The red blood cells of birds differ from mammalian red blood cells because:
1. they are white and have nuclei
2. they do not have nuclei
3. they have nuclei
4. they fight disease
Answer
C
Free Response
Describe the cause of different blood type groups.
Answer
Red blood cells are coated with proteins called antigens made of glycolipids and glycoproteins. When type A and type B blood are mixed, the blood agglutinates because of antibodies in the plasma that bind with the opposing antigen. Type O blood has no antigens. The Rh blood group has either the Rh antigen (Rh+) or no Rh antigen (Rh–).
List some of the functions of blood in the body.
Answer
Blood is important for regulation of the body’s pH, temperature, and osmotic pressure, the circulation of nutrients and removal of wastes, the distribution of hormones from endocrine glands, the elimination of excess heat; it also contains components for the clotting of blood to prevent blood loss. Blood also transports clotting factors and disease-fighting agents.
How does the lymphatic system work with blood flow?
Answer
Lymph capillaries take fluid from the blood to the lymph nodes. The lymph nodes filter the lymph by percolation through 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.
40.3: Mammalian Heart and Blood Vessels
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.
Review Questions
The heart’s internal pacemaker beats by:
1. an internal implant that sends an electrical impulse through the heart
2. the excitation of cardiac muscle cells at the sinoatrial node followed by the atrioventricular node
3. the excitation of cardiac muscle cells at the atrioventricular node followed by the sinoatrial node
4. the action of the sinus
Answer
B
During the systolic phase of the cardiac cycle, the heart is ________.
1. contracting
2. relaxing
3. contracting and relaxing
4. filling with blood
Answer
A
Cardiomyocytes are similar to skeletal muscle because:
1. they beat involuntarily
2. they are used for weight lifting
3. they pulse rhythmically
4. they are striated
Answer
D
How do arteries differ from veins?
1. Arteries have thicker smooth muscle layers to accommodate the changes in pressure from the heart.
2. Arteries carry blood.
3. Arteries have thinner smooth muscle layers and valves and move blood by the action of skeletal muscle.
4. Arteries are thin walled and are used for gas exchange.
Answer
A
Free Response
Describe the cardiac cycle.
Answer
The heart receives an electrical signal from the sinoatrial node triggering the cardiac muscle cells in the atria to contract. The signal pauses at the atrioventricular node before spreading to the walls of the ventricles so the blood is pumped through the body. This is the systolic phase. The heart then relaxes in the diastole and fills again with blood.
What happens in capillaries?
Answer
The capillaries basically exchange materials with their surroundings. Their walls are very thin and are made of one or two layers of cells, where gases, nutrients, and waste are diffused. They are distributed as beds, complex networks that link arteries as well as veins.
40.4: Blood Flow and Blood Pressure Regulation
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.
Review Questions
High blood pressure would be a result of ________.
1. a high cardiac output and high peripheral resistance
2. a high cardiac output and low peripheral resistance
3. a low cardiac output and high peripheral resistance
4. a low cardiac output and low peripheral resistance
Answer
A
Free Response
How does blood pressure change during heavy exercise?
Answer
The heart rate increases, which increases the hydrostatic pressure against the artery walls. At the same time, the arterioles dilate in response to the increased exercise, which reduces peripheral resistance. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/40%3A_The_Circulatory_System/40.E%3A_The_Circulatory_System_%28Exercises%29.txt |
To achieve a healthy balance, the human body should excrete the eight to ten glasses of water every day. This occurs via the processes of urination, defecation, sweating and, to a small extent, respiration. The organs and tissues of the human body are soaked in fluids that are maintained at constant temperature, pH, and solute concentration, all crucial elements of homeostasis. The solutes in body fluids are mainly mineral salts and sugars, and osmotic regulation is the process by which the mineral salts and water are kept in balance. Osmotic homeostasis is maintained despite the influence of external factors like temperature, diet, and weather conditions.
• 41.0: Prelude to Osmotic Regulation and Excretion
The daily intake recommendation for human water consumption is eight to ten glasses of water. In order to achieve a healthy balance, the human body should excrete the eight to ten glasses of water every day. This occurs via the processes of urination, defecation, sweating and, to a small extent, respiration.
• 41.1: Osmoregulation 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.
• 41.2: The Kidneys and 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.
• 41.3: Excretion Systems
Microorganisms and invertebrate animals use more primitive and simple mechanisms to get rid of their metabolic wastes than the mammalian system of kidney and urinary function. Three excretory systems evolved in organisms before complex kidneys: vacuoles, flame cells, and Malpighian tubules.
• 41.4: Nitrogenous Wastes
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.
• 41.5: 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.
• 41.E: Osmotic Regulation and Excretion (Exercises)
Thumbnail: Cross section of a kidney. (CC BY 3.0; Holly Fischer via Wikimedia Commons).
41: Osmotic Regulation and Excretion
The daily intake recommendation for human water consumption is eight to ten glasses of water. In order to achieve a healthy balance, the human body should excrete the eight to ten glasses of water every day. This occurs via the processes of urination, defecation, sweating and, to a small extent, respiration. The organs and tissues of the human body are soaked in fluids that are maintained at constant temperature, pH, and solute concentration, all crucial elements of homeostasis. The solutes in body fluids are mainly mineral salts and sugars, and osmotic regulation is the process by which the mineral salts and water are kept in balance. Osmotic homeostasis is maintained despite the influence of external factors like temperature, diet, and weather conditions. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.0%3A_Prelude_to_Osmotic_Regulation_and_Excretion.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.1%3A_Osmoregulation_and_Osmotic_Balance.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.2%3A_The_Kidneys_and_Osmoregulatory_Organs.txt |
Skills to Develop
• Explain how vacuoles, present in microorganisms, work to excrete waste
• Describe the way in which flame cells and nephridia in worms perform excretory functions and maintain osmotic balance
• Explain how insects use Malpighian tubules to excrete wastes and maintain osmotic balance
Microorganisms and invertebrate animals use more primitive and simple mechanisms to get rid of their metabolic wastes than the mammalian system of kidney and urinary function. Three excretory systems evolved in organisms before complex kidneys: vacuoles, flame cells, and Malpighian tubules.
Contractile Vacuoles in Microorganisms
The most fundamental feature of life is the presence of a cell. In other words, a cell is the simplest functional unit of a life. Bacteria are unicellular, prokaryotic organisms that have some of the least complex life processes in place; however, prokaryotes such as bacteria do not contain membrane-bound vacuoles. The cells of microorganisms like bacteria, protozoa, and fungi are bound by cell membranes and use them to interact with the environment. Some cells, including some leucocytes in humans, are able to engulf food by endocytosis—the formation of vesicles by involution of the cell membrane within the cells. The same vesicles are able to interact and exchange metabolites with the intracellular environment. In some unicellular eukaryotic organisms such as the amoeba, shown in Figure \(1\), cellular wastes and excess water are excreted by exocytosis, when the contractile vacuoles merge with the cell membrane and expel wastes into the environment. Contractile vacuoles (CV) should not be confused with vacuoles, which store food or water.
Flame Cells of Planaria and Nephridia of Worms
As multi-cellular systems evolved to have organ systems that divided the metabolic needs of the body, individual organs evolved to perform the excretory function. Planaria are flatworms that live in fresh water. Their excretory system consists of two tubules connected to a highly branched duct system. The cells in the tubules are called flame cells (or protonephridia) because they have a cluster of cilia that looks like a flickering flame when viewed under the microscope, as illustrated in Figure \(2\)a. The cilia propel waste matter down the tubules and out of the body through excretory pores that open on the body surface; cilia also draw water from the interstitial fluid, allowing for filtration. Any valuable metabolites are recovered by reabsorption. Flame cells are found in flatworms, including parasitic tapeworms and free-living planaria. They also maintain the organism’s osmotic balance.
Earthworms (annelids) have slightly more evolved excretory structures called nephridia, illustrated in Figure \(2\)b. A pair of nephridia is present on each segment of the earthworm. They are similar to flame cells in that they have a tubule with cilia. Excretion occurs through a pore called the nephridiopore. They are more evolved than the flame cells in that they have a system for tubular reabsorption by a capillary network before excretion.
Malpighian Tubules of Insects
Malpighian tubules are found lining the gut of some species of arthropods, such as the bee illustrated in Figure \(3\). They are usually found in pairs and the number of tubules varies with the species of insect. Malpighian tubules are convoluted, which increases their surface area, and they are lined with microvilli for reabsorption and maintenance of osmotic balance. Malpighian tubules work cooperatively with specialized glands in the wall of the rectum. Body fluids are not filtered as in the case of nephridia; urine is produced by tubular secretion mechanisms by the cells lining the Malpighian tubules that are bathed in hemolymph (a mixture of blood and interstitial fluid that is found in insects and other arthropods as well as most mollusks). Metabolic wastes like uric acid freely diffuse into the tubules. There are exchange pumps lining the tubules, which actively transport H+ ions into the cell and K+ or Na+ ions out; water passively follows to form urine. The secretion of ions alters the osmotic pressure which draws water, electrolytes, and nitrogenous waste (uric acid) into the tubules. Water and electrolytes are reabsorbed when these organisms are faced with low-water environments, and uric acid is excreted as a thick paste or powder. Not dissolving wastes in water helps these organisms to conserve water; this is especially important for life in dry environments.
Link to Learning
Visit this site to see a dissected cockroach, including a close-up look at its Malpighian tubules.
Summary
Many systems have evolved for excreting wastes that are simpler than the kidney and urinary systems of vertebrate animals. The simplest system is that of contractile vacuoles present in microorganisms. Flame cells and nephridia in worms perform excretory functions and maintain osmotic balance. Some insects have evolved Malpighian tubules to excrete wastes and maintain osmotic balance.
Glossary
flame cell
(also, protonephridia) excretory cell found in flatworms
Malpighian tubule
excretory tubules found in arthropods
microvilli
cellular processes that increase the surface area of cells
nephridia
excretory structures found in annelids
nephridiopore
pore found at the end of nephridia | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.3%3A_Excretion_Systems.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.4%3A_Nitrogenous_Wastes.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.5%3A_Hormonal_Control_of_Osmoregulatory_Functions.txt |
41.1: Osmoregulation and Osmotic Balance
Review Questions
When a dehydrated human patient needs to be given fluids intravenously, he or she is given:
1. water, which is hypotonic with respect to body fluids
2. saline at a concentration that is isotonic with respect to body fluids
3. glucose because it is a non-electrolyte
4. blood
Answer
B
The sodium ion is at the highest concentration in:
1. intracellular fluid
2. extracellular fluid
3. blood plasma
4. none of the above
Answer
B
Cells in a hypertonic solution tend to:
1. shrink due to water loss
2. swell due to water gain
3. stay the same size due to water moving into and out of the cell at the same rate
4. none of the above
Answer
A
Free Response
Why is excretion important in order to achieve osmotic balance?
Answer
Excretion allows an organism to rid itself of waste molecules that could be toxic if allowed to accumulate. It also allows the organism to keep the amount of water and dissolved solutes in balance.
Why do electrolyte ions move across membranes by active transport?
Answer
Electrolyte ions often require special mechanisms to cross the semi-permeable membranes in the body. Active transport is the movement against a concentration gradient.
41.2: The Kidneys and Osmoregulatory Organs
Review Questions
The macula densa is/are:
1. present in the renal medulla.
2. dense tissue present in the outer layer of the kidney.
3. cells present in the DCT and collecting tubules.
4. present in blood capillaries.
Answer
C
The osmolarity of body fluids is maintained at ________.
1. 100 mOsm
2. 300 mOsm
3. 1000 mOsm
4. it is not constantly maintained
Answer
B
The gland located at the top of the kidney is the ________ gland.
1. adrenal
2. pituitary
3. thyroid
4. thymus
Answer
A
Free Response
Why are the loop of Henle and vasa recta important for the formation of concentrated urine?
Answer
The loop of Henle is part of the renal tubule that loops into the renal medulla. In the loop of Henle, the filtrate exchanges solutes and water with the renal medulla and the vasa recta (the peritubular capillary network). The vasa recta acts as the countercurrent exchanger. The kidneys maintain the osmolality of the rest of the body at a constant 300 mOsm by concentrating the filtrate as it passes through the loop of Henle.
Describe the structure of the kidney.
Answer
Externally, the kidneys are surrounded by three layers. 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, which is the concave part of the “bean” shape.
41.3: Excretion Systems
Review Questions
Active transport of K+ in Malpighian tubules ensures that:
1. water follows K+ to make urine
2. osmotic balance is maintained between waste matter and bodily fluids
3. both a and b
4. neither a nor b
Answer
C
Contractile vacuoles in microorganisms:
1. exclusively perform an excretory function
2. can perform many functions, one of which is excretion of metabolic wastes
3. originate from the cell membrane
4. both b and c
Answer
D
Flame cells are primitive excretory organs found in ________.
1. arthropods
2. annelids
3. mammals
4. flatworms
Answer
D
Free Response
Why might specialized organs have evolved for excretion of wastes?
Answer
The removal of wastes, which could otherwise be toxic to an organism, is extremely important for survival. Having organs that specialize in this process and that operate separately from other organs provides a measure of safety for the organism.
Explain two different excretory systems other than the kidneys.
Answer
(1) Microorganisms engulf food by endocytosis—the formation of vacuoles by involution of the cell membrane within the cells. The same vacuoles interact and exchange metabolites with the intracellular environment. Cellular wastes are excreted by exocytosis when the vacuoles merge with the cell membrane and excrete wastes into the environment. (2) Flatworms have an excretory system that consists of two tubules. The cells in the tubules are called flame cells; they have a cluster of cilia that propel waste matter down the tubules and out of the body. (3) Annelids have nephridia which have a tubule with cilia. Excretion occurs through a pore called the nephridiopore. Annelids have a system for tubular reabsorption by a capillary network before excretion. (4) Malpighian tubules are found in some species of arthropods. They are usually found in pairs, and the number of tubules varies with the species of insect. Malpighian tubules are convoluted, which increases their surface area, and they are lined with microvilli for reabsorption and maintenance of osmotic balance. Metabolic wastes like uric acid freely diffuse into the tubules. Potassium ion pumps line the tubules, which actively transport out K+ ions, and water follows to form urine. Water and electrolytes are reabsorbed when these organisms are faced with low-water environments, and uric acid is excreted as a thick paste or powder. By not dissolving wastes in water, these organisms conserve water.
41.4: Nitrogenous Wastes
Review Questions
BUN is ________.
1. blood urea nitrogen
2. blood uric acid nitrogen
3. an indicator of blood volume
4. an indicator of blood pressure
Answer
A
Human beings accumulate ________ before excreting nitrogenous waste.
1. nitrogen
2. ammonia
3. urea
4. uric acid
Answer
C
Free Response
In terms of evolution, why might the urea cycle have evolved in organisms?
Answer
It is believed that 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.
Compare and contrast the formation of urea and uric acid.
Answer
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 urea cycle utilizes five intermediate steps, catalyzed by five different enzymes, to convert ammonia to urea. Birds, reptiles, and insects, on the other hand, convert toxic ammonia to uric acid instead of urea. Conversion of ammonia to uric acid requires more energy and is much more complex than conversion of ammonia to urea.
41.5: Hormonal Control of Osmoregulatory Functions
Review Questions
Renin is made by ________.
1. granular cells of the juxtaglomerular apparatus
2. the kidneys
3. the nephrons
4. All of the above.
Answer
A
Patients with Addison's disease ________.
1. retain water
2. retain salts
3. lose salts and water
4. have too much aldosterone
Answer
C
Which hormone elicits the “fight or flight” response?
1. epinephrine
2. mineralcorticoids
3. anti-diuretic hormone
4. thyroxine
Answer
A
Free Response
Describe how hormones regulate blood pressure, blood volume, and kidney function.
Answer
Hormones are small molecules that act as messengers within the body. Different regions of the nephron bear specialized cells, which have receptors to respond to chemical messengers and hormones. The hormones carry messages to the kidney. These 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.
How does the renin-angiotensin-aldosterone mechanism function? Why is it controlled by the kidneys?
Answer
The renin-angiotensin-aldosterone system acts through several steps to produce angiotensin II, which acts to stabilize blood pressure and volume. 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. ACE (angiotensin converting enzyme) converts angiotensin I to angiotensin II. Angiotensin II raises blood pressure by constricting blood vessels. It triggers the release of 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 from the hypothalamus, which leads to water retention. It acts directly on the nephrons and decreases GFR. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/41%3A_Osmotic_Regulation_and_Excretion/41.E%3A_Osmotic_Regulation_and_Excretion_%28Exercises%29.txt |
The environment consists of numerous pathogens, which are agents, usually microorganisms, that cause diseases in their hosts. A host is the organism that is invaded and often harmed by a pathogen. Pathogens include bacteria, protists, fungi and other infectious organisms. We are constantly exposed to pathogens in food and water, on surfaces, and in the air. Mammalian immune systems evolved for protection from such pathogens; they are composed of an extremely diverse array of specialized cells and soluble molecules that coordinate a rapid and flexible defense system capable of providing protection from a majority of these disease agents.
• 42.0: Prelude to the Immune System
Components of the immune system constantly search the body for signs of pathogens. When pathogens are found, immune factors are mobilized to the site of an infection. The immune factors identify the nature of the pathogen, strengthen the corresponding cells and molecules to combat it efficiently, and then halt the immune response after the infection is cleared to avoid unnecessary host cell damage.
• 42.1: Innate Immune Response
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.
• 42.2: Adaptive Immune Response
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.
• 42.3: Antibodies
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.
• 42.4: Disruptions in the Immune System
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.
• 42.E: The Immune System (Exercises)
Thumbnail: Monocytes, a type of white blood cell (Giemsa stained). (CC BY-SA 3.0; Dr Graham Beards via Wikimedia Commons).
42: The Immune System
The environment consists of numerous pathogens, which are agents, usually microorganisms, that cause diseases in their hosts. A host is the organism that is invaded and often harmed by a pathogen. Pathogens include bacteria, protists, fungi and other infectious organisms. We are constantly exposed to pathogens in food and water, on surfaces, and in the air. Mammalian immune systems evolved for protection from such pathogens; they are composed of an extremely diverse array of specialized cells and soluble molecules that coordinate a rapid and flexible defense system capable of providing protection from a majority of these disease agents.
Components of the immune system constantly search the body for signs of pathogens. When pathogens are found, immune factors are mobilized to the site of an infection. The immune factors identify the nature of the pathogen, strengthen the corresponding cells and molecules to combat it efficiently, and then halt the immune response after the infection is cleared to avoid unnecessary host cell damage. The immune system can remember pathogens to which it has been exposed to create a more efficient response upon re-exposure. This memory can last several decades. Features of the immune system, such as pathogen identification, specific response, amplification, retreat, and remembrance are essential for survival against pathogens. The immune response can be classified as either innate or active. The innate immune response is always present and attempts to defend against all pathogens rather than focusing on specific ones. Conversely, the adaptive immune response stores information about past infections and mounts pathogen-specific defenses.
Glossary
pathogen
an agent, usually a microorganism, that causes disease in the organisms that they invade
host
an organism that is invaded by a pathogen or parasite | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/42%3A_The_Immune_System/42.0%3A_Prelude_to_the_Immune_System.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/42%3A_The_Immune_System/42.1%3A_Innate_Immune_Response.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/42%3A_The_Immune_System/42.2%3A_Adaptive_Immune_Response.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/42%3A_The_Immune_System/42.3%3A_Antibodies.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/42%3A_The_Immune_System/42.4%3A_Disruptions_in_the_Immune_System.txt |
42.1: Innate Immune Response
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.
Review Questions
Which of the following is a barrier against pathogens provided by the skin?
1. high pH
2. mucus
3. tears
4. desiccation
Answer
D
Although interferons have several effects, they are particularly useful against infections with which type of pathogen?
1. bacteria
2. viruses
3. fungi
4. helminths
Answer
B
Which organelle do phagocytes use to digest engulfed particles?
1. lysosome
2. nucleus
3. endoplasmic reticulum
4. mitochondria
Answer
A
Which innate immune system component uses MHC I molecules directly in its defense strategy?
1. macrophages
2. neutrophils
3. NK cells
4. interferon
Answer
C
Free Response
Different MHC I molecules between donor and recipient cells can lead to rejection of a transplanted organ or tissue. Suggest a reason for this.
Answer
If the MHC I molecules expressed on donor cells differ from the MHC I molecules expressed on recipient cells, NK cells may identify the donor cells as “non-self” and produce perforin and granzymes to induce the donor cells to undergo apoptosis, which would destroy the transplanted organ.
If a series of genetic mutations prevented some, but not all, of the complement proteins from binding antibodies or pathogens, would the entire complement system be compromised?
Answer
The entire complement system would probably be affected even when only a few members were mutated such that they could no longer bind. Because the complement involves the binding of activated proteins in a specific sequence, when one or more proteins in the sequence are absent, the subsequent proteins would be incapable of binding to elicit the complement’s pathogen-destructive effects.
42.2: Adaptive Immune Response
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.
Review Questions
Which of the following is both a phagocyte and an antigen-presenting cell?
1. NK cell
2. eosinophil
3. neutrophil
4. macrophage
Answer
D
Which immune cells bind MHC molecules on APCs via CD8 coreceptors on their cell surfaces?
1. TH cells
2. CTLs
3. mast cells
4. basophils
Answer
B
What “self” pattern is identified by NK cells?
1. altered self
2. missing self
3. normal self
4. non-self
Answer
B
The acquired ability to prevent an unnecessary or destructive immune reaction to a harmless foreign particle, such as a food protein, is called ________.
1. the TH2 response
2. allergy
3. immune tolerance
4. autoimmunity
Answer
C
A memory B cell can differentiate upon re-exposure to a pathogen of which cell type?
1. CTL
2. naïve B cell
3. memory T cell
4. plasma cell
Answer
D
Foreign particles circulating in the blood are filtered by the ________.
1. spleen
2. lymph nodes
3. MALT
4. lymph
Answer
A
Free Response
Explain the difference between an epitope and an antigen.
Answer
An antigen is a molecule that reacts with some component of the immune response (antibody, B cell receptor, T cell receptor). An epitope is the region on the antigen through which binding with the immune component actually occurs.
What is a naïve B or T cell?
Answer
A naïve T or B cell is one that has not been activated by binding to the appropriate epitope. Naïve T and B cells cannot produce responses.
How does the TH1 response differ from the TH2 response?
Answer
The TH1 response involves the secretion of cytokines to stimulate macrophages and CTLs and improve their destruction of intracellular pathogens and tumor cells. It is associated with inflammation. The TH2 response is involved in the stimulation of B cells into plasma cells that synthesize and secrete antibodies.
In mammalian adaptive immune systems, T cell receptors are extraordinarily diverse. What function of the immune system results from this diversity, and how is this diversity achieved?
Answer
The diversity of TCRs allows the immune system to have millions of different T cells, and thereby to be specific in distinguishing antigens. This diversity arises from mutation and recombination in the genes that encode the variable regions of TCRs.
How do B and T cells differ with respect to antigens that they bind?
Answer
T cells bind antigens that have been digested and embedded in MHC molecules by APCs. In contrast, B cells function themselves as APCs to bind intact, unprocessed antigens.
Why is the immune response after reinfection much faster than the adaptive immune response after the initial infection?
Answer
Upon reinfection, the memory cells will immediately differentiate into plasma cells and CTLs without input from APCs or TH cells. In contrast, the adaptive immune response to the initial infection requires time for naïve B and T cells with the appropriate antigen specificities to be identified and activated.
42.3: Antibodies
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.
Review Questions
The structure of an antibody is similar to the extracellular component of which receptor?
1. MHC I
2. MHC II
3. BCR
4. none of the above
Answer
C
The first antibody class to appear in the serum in response to a newly encountered pathogen is ________.
1. IgM
2. IgA
3. IgG
4. IgE
Answer
A
What is the most abundant antibody class detected in the serum upon reexposure to a pathogen or in reaction to a vaccine?
1. IgM
2. IgA
3. IgG
4. IgE
Answer
C
Breastfed infants typically are resistant to disease because of ________.
1. active immunity
2. passive immunity
3. immune tolerance
4. immune memory
Answer
B
Free Response
What are the benefits and costs of antibody cross reactivity?
Answer
Cross reactivity of antibodies can be beneficial when it allows an individual's immune system to respond to an array of similar pathogens after being exposed to just one of them. A potential cost of cross reactivity is an antibody response to parts of the body (self) in addition to the appropriate antigen.
42.4: Disruptions in the Immune System
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.
Review Questions
Allergy to pollen is classified as:
1. an autoimmune reaction
2. immunodeficiency
3. delayed hypersensitivity
4. immediate hypersensitivity
Answer
D
A potential cause of acquired autoimmunity is ________.
1. tissue hypersensitivity
2. molecular mimicry
3. histamine release
4. radiation exposure
Answer
B
Autoantibodies are probably involved in:
1. reactions to poison ivy
2. pollen allergies
3. systemic lupus erythematosus
4. HIV/AIDS
Answer
C
Which of the following diseases is not due to autoimmunity?
1. rheumatic fever
2. systemic lupus erythematosus
3. diabetes mellitus
4. HIV/AIDS
Answer
D | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/42%3A_The_Immune_System/42.E%3A_The_Immune_System_%28Exercises%29.txt |
Animal reproduction is necessary for the survival of a species. In the animal kingdom, there are innumerable ways that species reproduce. Asexual reproduction produces genetically identical organisms (clones), whereas in sexual reproduction, the genetic material of two individuals combines to produce offspring that are genetically different from their parents.
• 43.0: Prelude to Animal Reproduction and Development
During sexual reproduction the male gamete (sperm) may be placed inside the female’s body for internal fertilization, or the sperm and eggs may be released into the environment for external fertilization. Seahorses provide an example of the latter. Following a mating dance, the female lays eggs in the male seahorse’s abdominal brood pouch where they are fertilized. The eggs hatch and the offspring develop in the pouch for several weeks.
• 43.1: Reproduction Methods
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.
• 43.2: Fertilization
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.
• 43.3: Human Reproductive Anatomy and Gametogenesis
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.
• 43.4: Hormonal Control of Human Reproduction
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.
• 43.5: Human Pregnancy and Birth
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.
• 43.6: Fertilization and Early Embryonic 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.
• 43.7: Organogenesis and Vertebrate Formation
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.
• 43.E: Animal Reproduction and Development (Exercises)
Thumbnail: Views of a Foetus in the Womb, detail. (Public Domain; Leonardo da Vinci via Wikimedia Commons)
43: Animal Reproduction and Development
Animal reproduction is necessary for the survival of a species. In the animal kingdom, there are innumerable ways that species reproduce. Asexual reproduction produces genetically identical organisms (clones), whereas in sexual reproduction, the genetic material of two individuals combines to produce offspring that are genetically different from their parents. During sexual reproduction the male gamete (sperm) may be placed inside the female’s body for internal fertilization, or the sperm and eggs may be released into the environment for external fertilization. Seahorses provide an example of the latter. Following a mating dance, the female lays eggs in the male seahorse’s abdominal brood pouch where they are fertilized. The eggs hatch and the offspring develop in the pouch for several weeks. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.0%3A_Prelude_to_Animal_Reproduction_and_Development.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.1%3A_Reproduction_Methods.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.2%3A_Fertilization.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 | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.3%3A_Human_Reproductive_Anatomy_and_Gametogenesis.txt |
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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.4%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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.5%3A_Human_Pregnancy_and_Birth.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.6%3A_Fertilization_and_Early_Embryonic_Development.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/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.7%3A_Organogenesis_and_Vertebrate_Formation.txt |
43.1: Reproduction Methods
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.
Review Questions
Which form of reproduction is thought to be best in a stable environment?
1. asexual
2. sexual
3. budding
4. parthenogenesis
Answer
A
Which form of reproduction can result from damage to the original animal?
1. asexual
2. fragmentation
3. budding
4. parthenogenesis
Answer
B
Which form of reproduction is useful to an animal with little mobility that reproduces sexually?
1. fission
2. budding
3. parthenogenesis
4. hermaphroditism
Answer
D
Genetically unique individuals are produced through ________.
1. sexual reproduction
2. parthenogenesis
3. budding
4. fragmentation
Answer
A
Free Response
Why is sexual reproduction useful if only half the animals can produce offspring and two separate cells must be combined to form a third?
Answer
Sexual reproduction produces a new combination of genes in the offspring that may better enable them to survive changes in the environment and assist in the survival of the species.
What determines which sex will result in offspring of birds and mammals?
Answer
The presence of the W chromosome in birds determines femaleness and the presence of the Y chromosome in mammals determines maleness. The absence of those chromosomes and the homogeneity of the offspring (ZZ or XX) leads to the development of the other sex.
43.2: Fertilization
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.
Review Questions
External fertilization occurs in which type of environment?
1. aquatic
2. forested
3. savanna
4. steppe
Answer
A
Which term applies to egg development within the female with nourishment derived from a yolk?
1. oviparity
2. viviparity
3. ovoviparity
4. ovovoparity
Answer
C
Which term applies to egg development outside the female with nourishment derived from a yolk?
1. oviparity
2. viviparity
3. ovoviparity
4. ovovoparity
Answer
A
Free Response
What are the advantages and disadvantages of external and internal forms of fertilization?
Answer
External fertilization can create large numbers of offspring without requiring specialized delivery or reproductive support organs. Offspring develop and mature quickly compared to internally fertilizing species. A disadvantage is that the offspring are out in the environment and predation can account for large loss of offspring. The embryos are susceptible to changes in the environment, which further depletes their numbers. Internally fertilizing species control their environment and protect their offspring from predators but must have specialized organs to complete these tasks and usually produce fewer embryos.
Why would paired external fertilization be preferable to group spawning?
Answer
Paired external fertilization allows the female to select the male for mating. It also has a greater chance of fertilization taking place, whereas spawning just puts a large number of sperm and eggs together and random interactions result in the fertilization.
43.3: Human Reproductive Anatomy and Gametogenesis
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.
Review Questions
Sperm are produced in the ________.
1. scrotum
2. seminal vesicles
3. seminiferous tubules
4. prostate gland
Answer
C
Most of the bulk of semen is made by the ________.
1. scrotum
2. seminal vesicles
3. seminiferous tubules
4. prostate gland
Answer
C
Which of the following cells in spermatogenesis is diploid?
1. primary spermatocyte
2. secondary spermatocyte
3. spermatid
4. sperm
Answer
A
Which female organ has the same embryonic origin as the penis?
1. clitoris
2. labia majora
3. greater vestibular glands
4. vagina
Answer
A
Which female organ has an endometrial lining that will support a developing baby?
1. labia minora
2. breast
3. ovaries
4. uterus
Answer
D
How many eggs are produced as a result of one meiotic series of cell divisions?
1. one
2. two
3. three
4. four
Answer
A
Free Response
Describe the phases of the human sexual response.
Answer
In phase one (excitement), vasodilation leads to vasocongestion and enlargement of erectile tissues. Vaginal secretions are released to lubricate the vagina during intercourse. In phase two (plateau), stimulation continues, the outer third of the vaginal wall enlarges with blood, and breathing and heart rate increase. In phase three (orgasm), rhythmic, involuntary contractions of muscles occur. In the male, reproductive accessory glands and tubules constrict, depositing 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. In phase four (resolution), the processes listed 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. Women do not experience a refractory period.
Compare spermatogenesis and oogenesis as to timing of the processes and the number and type of cells finally produced.
Answer
Stem cells are laid down in the male during gestation and lie dormant until adolescence. Stem cells in the female increase to one to two million and enter the first meiotic division and are arrested in prophase. At adolescence, spermatogenesis begins and continues until death, producing the maximum number of sperm with each meiotic division. Oogenesis continues again at adolescence in batches of oogonia with each menstrual cycle. These oogonia finish the first meiotic division, producing a primary oocyte with most of the cytoplasm and its contents, and a second cell called a polar body containing 23 chromosomes. The second meiotic division results in a secondary oocyte and a second oocyte. At ovulation, a mature haploid egg is released. If this egg is fertilized, it finishes the second meiotic division, including the chromosomes donated by the sperm in the finished cell. This is a diploid, fertilized egg.
43.4: Hormonal Control of Human Reproduction
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.
Review Questions
Which hormone causes Leydig cells to make testosterone?
1. FSH
2. LH
3. inhibin
4. estrogen
Answer
A
Which hormone causes FSH and LH to be released?
1. testosterone
2. estrogen
3. GnRH
4. progesterone
Answer
C
Which hormone signals ovulation?
1. FSH
2. LH
3. inhibin
4. estrogen
Answer
B
Which hormone causes the re-growth of the endometrial lining of the uterus?
1. testosterone
2. estrogen
3. GnRH
4. progesterone
Answer
D
Free Response
If male reproductive pathways are not cyclical, how are they controlled?
Answer
Negative feedback in the male system is supplied through two hormones: inhibin and testosterone. Inhibin is produced by Sertoli cells when the sperm count exceeds set limits. The hormone inhibits GnRH and FSH, decreasing the activity of the Sertoli cells. Increased levels of testosterone affect the release of both GnRH and LH, decreasing the activity of the Leydig cells, resulting in decreased testosterone and sperm production.
Describe the events in the ovarian cycle leading up to ovulation.
Answer
Low levels of progesterone allow the hypothalamus to send GnRH to the anterior pituitary and cause the release of FSH and LH. FSH stimulates follicles on the ovary to grow and prepare the eggs for ovulation. As the follicles increase in size, they begin to release estrogen and a low level of progesterone into the blood. The level of estrogen rises to a peak, causing a spike in the concentration of LH. This causes the most mature follicle to rupture and ovulation occurs.
43.5: Human Pregnancy and Birth
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.
Review Questions
Nutrient and waste requirements for the developing fetus are handled during the first few weeks by:
1. the placenta
2. diffusion through the endometrium
3. the chorion
4. the blastocyst
Answer
B
Progesterone is made during the third trimester by the:
1. placenta
2. endometrial lining
3. chorion
4. corpus luteum
Answer
A
Which contraceptive method is 100 percent effective at preventing pregnancy?
1. condom
2. oral hormonal methods
3. sterilization
4. abstinence
Answer
D
Which type of short term contraceptive method is generally more effective than others?
1. barrier
2. hormonal
3. natural family planning
4. withdrawal
Answer
B
Which hormone is primarily responsible for the contractions during labor?
1. oxytocin
2. estrogen
3. β-HCG
4. progesterone
Answer
A
Major organs begin to develop during which part of human gestation?
1. fertilization
2. first trimester
3. second trimester
4. third trimester
Answer
B
Free Response
Describe the major developments during each trimester of human gestation.
Answer
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 established during the first trimester. The placenta takes over the production of estrogen and high levels of progesterone and handles the nutrient and waste requirements of the fetus. The third trimester exhibits the greatest growth of the fetus, culminating in labor and delivery.
Describe the stages of labor.
Answer
Stage one of labor results in the thinning of the cervix and the dilation of the cervical opening. Stage two delivers the baby, and stage three delivers the placenta.
43.6: Fertilization and Early Embryonic 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.
Review Questions
Which of the following is false?
1. The endoderm, mesoderm, ectoderm are germ layers.
2. The trophoblast is a germ layer.
3. The inner cell mass is a source of embryonic stem cells.
4. The blastula is often a hollow ball of cells.
Answer
B
During cleavage, the mass of cells:
1. increases
2. decreases
3. doubles with every cell division
4. does not change significantly
Answer
D
Free Response
What do you think would happen if multiple sperm fused with one egg?
Answer
Multiple sperm can fuse with the egg, resulting in polyspermy. The resulting embryo is not genetically viable and dies within a few days.
Why do mammalian eggs have a small concentration of yolk, while bird and reptile eggs have a large concentration of yolk?
Answer
Mammalian eggs do not need a lot of yolk because the developing fetus obtains nutrients from the mother. Other species, in which the fetus develops outside of the mother’s body, such as occurs with birds, require a lot of yolk in the egg to nourish the embryo during development.
43.7: Organogenesis and Vertebrate Formation
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.
Review Questions
Which of the following gives rise to the skin cells?
1. ectoderm
2. endoderm
3. mesoderm
4. none of the above
Answer
A
The ribs form from the ________.
1. notochord
2. neural plate
3. neural tube
4. somites
Answer
D
Free Response
Explain how the different germ layers give rise to different tissue types.
Answer
Organs form from the germ layers through the process of differentiation. During differentiation, the embryonic stem cells express a specific set of genes that will determine their ultimate fate as a 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.
Explain the role of axis formation in development.
Answer
Animal bodies have lateral-medial (left-right), dorsal-ventral (back-belly), and anterior-posterior (head-feet) axes. The dorsal cells are genetically programmed to form the notochord and define the axis. There are many genes responsible for axis formation. Mutations in these genes lead to the loss of symmetry required for organism development. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/7%3A_Animal_Structure_and_Function/43%3A_Animal_Reproduction_and_Development/43.E%3A_Animal_Reproduction_and_Development_%28Exercises%29.txt |
Humans are a part of the ecological landscape, and human health is one important part of human interaction with our physical and living environment.
• 44.0: Prelude to Ecology and the Biosphere
Why study ecology? Perhaps you are interested in learning about the natural world and how living things have adapted to the physical conditions of their environment. Or, perhaps you’re a future physician seeking to understand the connection between human health and ecology.
• 44.1: The Scope of Ecology
Ecology is the study of the interactions of living organisms with their environment. One core goal of ecology is to understand the distribution and abundance of living things in the physical environment. Attainment of this goal requires the integration of scientific disciplines inside and outside of biology, such as biochemistry, physiology, evolution, biodiversity, molecular biology, geology, and climatology.
• 44.2: Biogeography
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 the biosphere.
• 44.3: 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 biome distribution.
• 44.4: 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.
• 44.5: Climate and the Effects of Global Climate Change
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.
• 44.E: Ecology and the Biosphere (Exercises)
44: Ecology and the Biosphere
Why study ecology? Perhaps you are interested in learning about the natural world and how living things have adapted to the physical conditions of their environment. Or, perhaps you’re a future physician seeking to understand the connection between human health and ecology.
Humans are a part of the ecological landscape, and human health is one important part of human interaction with our physical and living environment. Lyme disease, for instance, serves as one modern-day example of the connection between our health and the natural world. More formally known as Lyme borreliosis, Lyme disease is a bacterial infection that can be transmitted to humans when they are bitten by the deer tick (Ixodes scapularis), which is the primary vector for this disease. However, not all deer ticks carry the bacteria that will cause Lyme disease in humans, and I. scapularis can have other hosts besides deer. In fact, it turns out that the probability of infection depends on the type of host upon which the tick develops: a higher proportion of ticks that live on white-footed mice carry the bacterium than do ticks that live on deer. Knowledge about the environments and population densities in which the host species is abundant would help a physician or an epidemiologist better understand how Lyme disease is transmitted and how its incidence could be reduced. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.0%3A_Prelude_to_Ecology_and_the_Biosphere.txt |
Skills to Develop
• Define ecology and the four levels of ecological research
• Describe examples of the ways in which ecology requires the integration of different scientific disciplines
• Distinguish between abiotic and biotic components of the environment
• Recognize the relationship between abiotic and biotic components of the environment
Ecology is the study of the interactions of living organisms with their environment. One core goal of ecology is to understand the distribution and abundance of living things in the physical environment. Attainment of this goal requires the integration of scientific disciplines inside and outside of biology, such as biochemistry, physiology, evolution, biodiversity, molecular biology, geology, and climatology. Some ecological research also applies aspects of chemistry and physics, and it frequently uses mathematical models.
Link to Learning
Climate change can alter where organisms live, which can sometimes directly affect human health. Watch the PBS video “Feeling the Effects of Climate Change” in which researchers discover a pathogenic organism living far outside of its normal range.
Levels of Ecological Study
When a discipline such as biology is studied, it is often helpful to subdivide it into smaller, related areas. For instance, cell biologists interested in cell signaling need to understand the chemistry of the signal molecules (which are usually proteins) as well as the result of cell signaling. Ecologists interested in the factors that influence the survival of an endangered species might use mathematical models to predict how current conservation efforts affect endangered organisms. To produce a sound set of management options, a conservation biologist needs to collect accurate data, including current population size, factors affecting reproduction (like physiology and behavior), habitat requirements (such as plants and soils), and potential human influences on the endangered population and its habitat (which might be derived through studies in sociology and urban ecology). Within the discipline of ecology, researchers work at four specific levels, sometimes discretely and sometimes with overlap: organism, population, community, and ecosystem (Figure \(1\)).
Organismal Ecology
Researchers studying ecology at the organismal level are interested in the adaptations that enable individuals to live in specific habitats. These adaptations can be morphological, physiological, and behavioral. For instance, the Karner blue butterfly (Lycaeides melissa samuelis) (Figure \(2\)) is considered a specialist because the females preferentially oviposit (that is, lay eggs) on wild lupine. This preferential adaptation means that the Karner blue butterfly is highly dependent on the presence of wild lupine plants for its continued survival.
After hatching, the larval caterpillars emerge and spend four to six weeks feeding solely on wild lupine (Figure \(3\)). The caterpillars pupate (undergo metamorphosis) and emerge as butterflies after about four weeks. The adult butterflies feed on the nectar of flowers of wild lupine and other plant species. A researcher interested in studying Karner blue butterflies at the organismal level might, in addition to asking questions about egg laying, ask questions about the butterflies’ preferred temperature (a physiological question) or the behavior of the caterpillars when they are at different larval stages (a behavioral question).
Population Ecology
A population is a group of interbreeding organisms that are members of the same species living in the same area at the same time. (Organisms that are all members of the same species are called conspecifics.) A population is identified, in part, by where it lives, and its area of population may have natural or artificial boundaries: natural boundaries might be rivers, mountains, or deserts, while examples of artificial boundaries include mowed grass, manmade structures, or roads. The study of population ecology focuses on the number of individuals in an area and how and why population size changes over time. Population ecologists are particularly interested in counting the Karner blue butterfly, for example, because it is classified as federally endangered. However, the distribution and density of this species is highly influenced by the distribution and abundance of wild lupine. Researchers might ask questions about the factors leading to the decline of wild lupine and how these affect Karner blue butterflies. For example, ecologists know that wild lupine thrives in open areas where trees and shrubs are largely absent. In natural settings, intermittent wildfires regularly remove trees and shrubs, helping to maintain the open areas that wild lupine requires. Mathematical models can be used to understand how wildfire suppression by humans has led to the decline of this important plant for the Karner blue butterfly.
Community Ecology
A biological community consists of the different species within an area, typically a three-dimensional space, and the interactions within and among these species. Community ecologists are interested in the processes driving these interactions and their consequences. Questions about conspecific interactions often focus on competition among members of the same species for a limited resource. Ecologists also study interactions among various species; members of different species are called heterospecifics. Examples of heterospecific interactions include predation, parasitism, herbivory, competition, and pollination. These interactions can have regulating effects on population sizes and can impact ecological and evolutionary processes affecting diversity.
For example, Karner blue butterfly larvae form mutualistic relationships with ants. Mutualism is a form of a long-term relationship that has coevolved between two species and from which each species benefits. For mutualism to exist between individual organisms, each species must receive some benefit from the other as a consequence of the relationship. Researchers have shown that there is an increase in the probability of survival when Karner blue butterfly larvae (caterpillars) are tended by ants. This might be because the larvae spend less time in each life stage when tended by ants, which provides an advantage for the larvae. Meanwhile, the Karner blue butterfly larvae secrete a carbohydrate-rich substance that is an important energy source for the ants. Both the Karner blue larvae and the ants benefit from their interaction.
Ecosystem Ecology
Ecosystem ecology is an extension of organismal, population, and community ecology. The ecosystem is composed of all the biotic components (living things) in an area along with the abiotic components (non-living things) of that area. Some of the abiotic components include air, water, and soil. Ecosystem biologists ask questions about how nutrients and energy are stored and how they move among organisms and the surrounding atmosphere, soil, and water.
The Karner blue butterflies and the wild lupine live in an oak-pine barren habitat. This habitat is characterized by natural disturbance and nutrient-poor soils that are low in nitrogen. The availability of nutrients is an important factor in the distribution of the plants that live in this habitat. Researchers interested in ecosystem ecology could ask questions about the importance of limited resources and the movement of resources, such as nutrients, though the biotic and abiotic portions of the ecosystem.
Career Connection: Ecologist
A career in ecology contributes to many facets of human society. Understanding ecological issues can help society meet the basic human needs of food, shelter, and health care. Ecologists can conduct their research in the laboratory and outside in natural environments (Figure \(4\)). These natural environments can be as close to home as the stream running through your campus or as far away as the hydrothermal vents at the bottom of the Pacific Ocean. Ecologists manage natural resources such as white-tailed deer populations (Odocoileus virginianus) for hunting or aspen (Populus spp.) timber stands for paper production. Ecologists also work as educators who teach children and adults at various institutions including universities, high schools, museums, and nature centers. Ecologists may also work in advisory positions assisting local, state, and federal policymakers to develop laws that are ecologically sound, or they may develop those policies and legislation themselves. To become an ecologist requires an undergraduate degree, usually in a natural science. The undergraduate degree is often followed by specialized training or an advanced degree, depending on the area of ecology selected. Ecologists should also have a broad background in the physical sciences, as well as a sound foundation in mathematics and statistics.
Link to Learning
Visit this site to see Stephen Wing, a marine ecologist from the University of Otago, discuss the role of an ecologist and the types of issues ecologists explore.
Summary
Ecology is the study of the interactions of living things with their environment. Ecologists ask questions across four levels of biological organization—organismal, population, community, and ecosystem. At the organismal level, ecologists study individual organisms and how they interact with their environments. At the population and community levels, ecologists explore, respectively, how a population of organisms changes over time and the ways in which that population interacts with other species in the community. Ecologists studying an ecosystem examine the living species (the biotic components) of the ecosystem as well as the nonliving portions (the abiotic components), such as air, water, and soil, of the environment.
Glossary
abiotic
nonliving components of the environment
biotic
living components of the environment
conspecifics
individuals that are members of the same species
ecology
study of interaction between living things and their environment
heterospecifics
individuals that are members of different species | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.1%3A_The_Scope_of_Ecology.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/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.2%3A_Biogeography.txt |
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/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.3%3A_Terrestrial_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/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.4%3A_Aquatic_Biomes.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/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.5%3A_Climate_and_the_Effects_of_Global_Climate_Change.txt |
44.1: The Scope of Ecology
Ecology is the study of the interactions of living organisms with their environment. One core goal of ecology is to understand the distribution and abundance of living things in the physical environment. Attainment of this goal requires the integration of scientific disciplines inside and outside of biology, such as biochemistry, physiology, evolution, biodiversity, molecular biology, geology, and climatology.
Review Questions
Which of the following is a biotic factor?
1. wind
2. disease-causing microbe
3. temperature
4. soil particle size
Answer
B
The study of nutrient cycling though the environment is an example of which of the following?
1. organismal ecology
2. population ecology
3. community ecology
4. ecosystem ecology
Answer
D
Free Response
Ecologists often collaborate with other researchers interested in ecological questions. Describe the levels of ecology that would be easier for collaboration because of the similarities of questions asked. What levels of ecology might be more difficult for collaboration?
Answer
Ecologists working in organismal or population ecology might ask similar questions about how the biotic and abiotic conditions affect particular organisms and, thus, might find collaboration to be mutually beneficial. Levels of ecology such as community ecology or ecosystem ecology might pose greater challenges for collaboration because these areas are very broad and may include many different environmental components.
The population is an important unit in ecology as well as other biological sciences. How is a population defined, and what are the strengths and weaknesses of this definition? Are there some species that at certain times or places are not in populations?
Answer
It is beneficial to consider a population to be all of the individuals living in the same area at the same time because it allows the ecologist to identify and study all of the abiotic and biotic factors that may affect the members of the population. However, this definition of a population could be considered a drawback if it prohibits the ecologist from studying a population’s individuals that may be transitory, but still influential. Some species with members that have a wide geographic range might not be considered to be a population, but could still have many of the qualities of a population.
44.2: Biogeography
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 the biosphere.
Review Questions
Understory plants in a temperate forest have adaptations to capture limited ________.
1. water
2. nutrients
3. heat
4. sunlight
Answer
D
An ecologist hiking up a mountain may notice different biomes along the way due to changes in all of the following except:
1. elevation
2. rainfall
3. latitude
4. temperature
Answer
C
Free Response
Compare and contrast ocean upwelling and spring and fall turnovers.
Answer
Ocean upwelling is a continual process that occurs year-round. Spring and fall turnover in freshwater lakes and ponds, however, is a seasonal process that occurs due to temperature changes in the water that take place during springtime warming and autumn cooling. Both ocean upwelling and spring and fall turnover enable nutrients in the organic materials at the bottom of the body of water to be recycled and reused by living things.
Many endemic species are found in areas that are geographically isolated. Suggest a plausible scientific explanation for why this is so.
Answer
Areas that have been geographically isolated for very long periods of time allow unique species to evolve; these species are distinctly different from those of surrounding areas and remain so, since geographic isolation keeps them separated from other species.
44.3: 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
Review Questions
Which of the following biomes is characterized by abundant water resources?
1. deserts
2. boreal forests
3. savannas
4. tropical wet forests
Answer
D
Which of the following biomes is characterized by short growing seasons?
1. deserts
2. tropical wet forests
3. Arctic tundras
4. savannas
Answer
C
Free Response
The extremely low precipitation of subtropical desert biomes might lead one to expect fire to be a major disturbance factor; however, fire is more common in the temperate grassland biome than in the subtropic desert biome. Why is this?
Answer
Fire is less common in desert biomes than in temperate grasslands because deserts have low net primary productivity and, thus, very little plant biomass to fuel a fire.
In what ways are the subtropical desert and the arctic tundra similar?
Answer
Both the subtropical desert and the arctic tundra have a low supply of water. In the desert, this is due to extremely low precipitation, and in the arctic tundra, much of the water is unavailable to plants because it is frozen. Both the subtropical desert and the arctic tundra have low net primary productivity.
44.4: 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.
Review Questions
Where would you expect to find the most photosynthesis in an ocean biome?
1. aphotic zone
2. abyssal zone
3. benthic realm
4. intertidal zone
Answer
D
A key feature of estuaries is:
1. low light conditions and high productivity
2. salt water and fresh water
3. frequent algal blooms
4. little or no vegetation
Answer
B
Free Response
Scientists have discovered the bodies of humans and other living things buried in bogs for hundreds of years, but not yet decomposed. Suggest a possible biological explanation for why such bodies are so well-preserved.
Answer
Bogs are low in oxygen and high in organic acids. The low oxygen content and the low pH both slow the rate of decomposition.
Describe the conditions and challenges facing organisms living in the intertidal zone.
Answer
Organisms living in the intertidal zone must tolerate periodic exposure to air and sunlight and must be able to be periodically dry. They also must be able to endure the pounding waves; for this reason, some shoreline organisms have hard exoskeletons that provide protection while also reducing the likelihood of drying out.
44.5: Climate and the Effects of Global Climate Change
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.
Review Questions
Which of the following is an example of a weather event?
1. The hurricane season lasts from June 1 through November 30.
2. The amount of atmospheric CO2 has steadily increased during the last century.
3. A windstorm blew down trees in the Boundary Waters Canoe Area in Minnesota on July 4, 1999.
4. Deserts are generally dry ecosystems having very little rainfall.
Answer
C
Which of the following natural forces is responsible for the release of carbon dioxide and other atmospheric gases?
1. the Milankovitch cycles
2. volcanoes
3. solar intensity
4. burning of fossil fuels
Answer
B
Free Response
Compare and contrast how natural- and human-induced processes have influenced global climate change.
Answer
Natural processes such as the Milankovitch cycles, variation in solar intensity, and volcanic eruptions can cause periodic, intermittent changes in global climate. Human activity, in the form of emissions from the burning of fossil fuels, has caused a progressive rise in the levels of atmospheric carbon dioxide.
Predict possible consequences if carbon emissions from fossil fuels continue to rise.
Answer
If carbon emissions continue to rise, the global temperature will continue to rise; thus, ocean waters will cause the rising of sea levels at the coastlines. Continued melting of glaciers and reduced spring and summer meltwaters may cause summertime water shortages. Changes in seasonal temperatures may alter lifecycles and interrupt breeding patterns in many species of plants and animals. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/8%3A_Ecology/44%3A_Ecology_and_the_Biosphere/44.E%3A_Ecology_and_the_Biosphere_%28Exercises%29.txt |
Community ecology is the study of the interactions between species in communities on many spatial and temporal scales, including the distribution, structure, abundance, demography, and interactions between coexisting populations. The primary focus of community ecology is on the interactions between populations as determined by specific genotypic and phenotypic characteristics.
• 45.0: Prelude to Population and Community Ecology
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.
• 45.1: Population Demography
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.
• 45.2: Life Histories and Natural Selection
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.
• 45.3: 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.
• 45.4: Population Dynamics and Regulation
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.
• 45.5: 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.
• 45.6: Community Ecology
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.
• 45.7: Behavioral Biology - Proximate and Ultimate Causes of Behavior
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.
• 45.E: Population and Community Ecology (Exercises)
45: Population and Community Ecology
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. | textbooks/bio/Introductory_and_General_Biology/General_Biology_1e_(OpenStax)/8%3A_Ecology/45%3A_Population_and_Community_Ecology/45.0%3A_Prelude_to_Population_and_Community_Ecology.txt |
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/General_Biology_1e_(OpenStax)/8%3A_Ecology/45%3A_Population_and_Community_Ecology/45.1%3A_Population_Demography.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/General_Biology_1e_(OpenStax)/8%3A_Ecology/45%3A_Population_and_Community_Ecology/45.2%3A_Life_Histories_and_Natural_Selection.txt |
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