Miller Harley-431-470

Miller−Harley: Zoology, III. Form and Function: A 27.
Nutrition and Digestion © The McGraw−Hill

Fifth Edition Comparative Perspecitive Companies, 2001
CHAPTER 27
NUTRITION AND DIGESTION
Outline Concepts
Evolution of Nutrition 1. Animals are heterotrophic organisms that use food to supply both raw materials and
The Metabolic Fates of Nutrients in energy. Nutrition includes all of those processes by which an animal takes in, digests,
Heterotrophs absorbs, stores, and uses food to meet its metabolic needs.
Calories and Energy 2. Digestion is the chemical and/or mechanical breakdown of food into particles that indi-
Macronutrients vidual cells of an organism can absorb. Digestion can occur either inside a cell (intracel-
Micronutrients lular), outside a cell (extracellular), or in both places.
Digestion 3. Most animals must work for their nutrients. The number of specializations that have
Animal Strategies for Getting and Using Food evolved for food procurement (feeding) and extracellular digestion are almost as numer-
Continuous Versus Discontinuous Feeders ous as the number of animal species. Some examples include continuous versus discon-
Suspension Feeders tinuous feeding, suspension feeding, deposit feeding, herbivory, predation, surface
Deposit Feeders
nutrient absorption, and fluid feeding.
Herbivory
4. Intracellular digestion occurs in some invertebrates (e.g., in sponges); others (e.g., some
Predation
cnidarians and molluscs) utilize both intracellular and extracellular digestion; and most
Surface Nutrient Absorption
higher invertebrates (e.g., insects) have evolved variations in extracellular digestion that
Fluid Feeders
allow them to exploit different food sources.
Diversity in Digestive Structures: Invertebrates
Protozoa
5. In primitive, multicellular animals, such as cnidarians, the gut is a blind (closed) sac
Bivalve Molluscs called a gastrovascular cavity. Its one opening is both entrance and exit; thus, it is an
Insects incomplete digestive tract. The development of an anus and complete digestive tract
Diversity in Digestive Structures: Vertebrates in the aschelminths was an evolutionary breakthrough. The many variations of the
Tongues basic complete digestive tract correlate with different food-gathering mechanisms
Teeth and diets.
Salivary Glands 6. Vertebrate digestive systems have evolved into assembly lines where food is first broken
Esophagi down mechanically and then chemically by digestive enzymes. The simple sugars, fats,
Stomachs triglycerides, amino acids, vitamins, and minerals that result are then taken into the cir-
Gizzards culatory systems for distribution throughout the animal’s body and are used in mainte-
Rumens nance, growth, and energy production.
Ceca
Livers and Gallbladders
Pancreata
Nutrition includes all of those processes by which an animal takes in, digests, absorbs,
Intestines
stores, and uses food (nutrients) to meet its metabolic needs. Digestion (L. digestio, from
The Mammalian Digestive System
⫹ dis, apart ⫹ gerere, to carry) is the chemical and/or mechanical breakdown of food into
Gastrointestinal Motility and Its Control
particles that individual cells of an animal can absorb. This chapter discusses animal nutri-
Oral Cavity
tion, the different strategies animals use for consuming and using food, and various animal
Pharynx and Esophagus
digestive systems.
Stomach
Small Intestine: Main Site of Digestion
Large Intestine
Role of the Pancreas in Digestion
Role of the Liver and Gallbladder in
Digestion This chapter contains evolutionary concepts, which are set off in this font.
433
Miller−Harley: Zoology, III. Form and Function: A 27. Nutrition and Digestion © The McGraw−Hill
434 PART THREE Form and Function: A Comparative Perspective
EVOLUTION OF NUTRITION MACRONUTRIENTS

Nutrients in the food an animal consumes provide the necessary With a few notable exceptions, heterotrophs require organic mol-
chemicals for growth, maintenance, and energy production. Over- ecules, such as carbohydrates, lipids, and proteins, in their diets.
all, the nutritional requirements of an animal are inversely related Enzymes break down these molecules into components that can
to its ability to synthesize molecules essential for life. The fewer be used for energy production or as sources for the “building
such biosynthetic abilities an animal has, the more kinds of nutri- blocks’’ of life.
ents it must obtain from its environment. Green plants and pho-
tosynthetic protists have the fewest such nutritional requirements Carbohydrates: Carbon and Energy
because they can synthesize all their own complex molecules from from Sugars and Starches
simpler inorganic substances; they are autotrophs (Gr. auto, self ⫹
trophe, nourishing). Animals, fungi, and bacteria that cannot syn- The major dietary source of energy for heterotrophs is complex
thesize many of their own organic molecules and must obtain carbohydrates (figure 27.1a). Most carbohydrates originally come
them by consuming other organisms or their products are het- from plant sources. Various polysaccharides, disaccharides, or any
erotrophs (Gr. heteros, another or different ⫹ trophe, nourishing). of a variety of simple sugars (monosaccharides) can meet this
Animals such as rabbits that subsist entirely on plant material are dietary need. Carbohydrates also are a major carbon source for
herbivores (L. herba, plant ⫹ vorare, to eat). Carnivores (L. caro, incorporation into important organic compounds. Many plants
flesh), such as hawks, are animals that eat only meat. Omnivores also supply cellulose, a polysaccharide that humans and other ani-
(L. omnius, all), such as humans, bears, raccoons, and pigs, eat mals (with the exception of herbivores) cannot digest. Cellulose is
both plant and animal matter. Insectivores, such as bats, eat pri- sometimes called dietary fiber. It assists in the passage of food
marily arthropods. through the alimentary canal of mammals. Cellulose may also
Losses of biosynthetic abilities have marked much of ani- reduce the risk of cancer of the colon, because the mutagenic
mal evolution. Once an animal routinely obtains essential, com- compounds that form during the storage of feces are reduced if
plex organic molecules in its diet, it can afford to lose the abil- fecal elimination is more frequent.
ity to synthesize those molecules. Moreover, the loss of this
ability confers a selective advantage on the animal because the Lipids: Highly Compact Energy-
animal stops expending energy and resources to synthesize Storage Nutrients
molecules that are already in its diet. Thus, as the diet of ani- Neutral lipids (fats) or triacylglycerols are contained in fats and
mals became more varied, they tended to lose their abilities to oils, meat and dairy products, nuts, and some fruits and vegetables
synthesize such widely available molecules as some of the high in fats, such as avocados (figure 27.1b). Lipids are the most
amino acids. concentrated source of food energy. They produce about 9 Calories
(kcal) of usable energy per gram, more than twice the energy avail-
able from an equal mass of carbohydrate or protein (table 27.1).
THE METABOLIC FATES OF Many heterotrophs have an absolute dietary requirement for
NUTRIENTS IN HETEROTROPHS lipids, sometimes for specific types. For example, many animals
require unsaturated fatty acids (e.g., linoleic acid, linolenic acid,
The nutrients that a heterotroph ingests can be divided into
and arachidonic acid). These fatty acids act as precursor molecules
macronutrients and micronutrients. Macronutrients are needed
for the synthesis of sterols, the most common of which is choles-
in large quantities and include the carbohydrates, lipids, and pro-
terol. The sterols are also required for the synthesis of steroid hor-
teins. Micronutrients are needed in small quantities and include
mones and cholesterol, which is incorporated into cell mem-
organic vitamins and inorganic minerals. Together, these nutri-
branes. Other lipids insulate the bodies of some vertebrates and
ents make up the animal’s dietary requirements. Besides these
help maintain a constant temperature.
nutrients, animals require water.
Proteins: Basic to the Structure and
CALORIES AND ENERGY Function of Cells
The energy value of food is measured in terms of calories or Calo- Animal sources of protein include other animals and milk. Plant
ries. A calorie (L. calor, heat) is the amount of energy required to sources include beans, peas, and nuts. Proteins are needed for their
raise the temperature of 1 g of water 1° C. A calorie, with a small amino acids, which heterotrophs use to build their own body pro-
c, is also called a gram calorie. A kilocalorie, also known as a teins (figure 27.1c).
Calorie or kilogram calorie (kcal), is equal to 1,000 calories. In
popular usage, you talk about calories but actually mean Calories, MICRONUTRIENTS
because the larger unit is more useful for measuring the energy
value of food. If an advertisement says that a so-called light beer Micronutrients are usually small ions, organic vitamins, inorganic
contains 95 calories per 12 oz, it really means 95,000 calories, 95 minerals, and molecules that are used repeatedly in enzymatic
Calories, or 95 kcal. reactions or as parts of certain proteins (e.g., copper in hemocyanin
CHAPTER 27 Nutrition and Digestion 435
(a) (b) (c)
Carbohydrates Lipids Proteins
Neutral
Sugar Starches Glycerol Steroids Amino acids
fats
Glucose Steroids the Proteins the

Fatty acids
animal needs animal needs
Other lipids the Nitrogen

Glycolysis
animal needs waste
Acetyl-CoA
Carbon Krebs
Electron
compounds cycle
transport
chain
ATP
energy
FIGURE 27.1
Macronutrients in the Diet. (a) Carbohydrate foods break down to their constituent sugars and starches, and ultimately into glucose. Individual cells
use this sugar in glycolysis and aerobic respiration to create new carbon compounds or ATP energy. (b) Lipids (fats and oils) in the diet break down to
neutral fats, glycerol, and steroids. These molecules can be modified and incorporated into the lipids or steroids the animal needs for storing fat or gener-
ating hormones, or they can be converted to acetyl-CoA and enter the Krebs cycle and electron transport chain for ATP production. (c) Proteins break
down to amino acids, which are incorporated into new proteins or modified to enter the Krebs cycle and electron transport chain to produce ATP energy.
nerve and muscle in an animal’s body. Animals lose large

TA B L E 2 7 . 1 quantities of these minerals, especially sodium, in the urine
THE AVERAGE CALORIC VALUES OF every day. Animals that sweat to help regulate body tempera-
MACRONUTRIENTS ture lose sodium in their sweat. A daily supply of calcium is
needed for muscular activity and, with phosphorus, for bone
MACRONUTRIENT CALORIES PER GRAM
formation. Table 27.2 lists the functions of the major essential
Carbohydrates 4.1 minerals.
Lipids 9.3 Other minerals are known as trace minerals, trace elements,
Proteins 4.4 or microminerals. Animals need these in only very small amounts
for various enzymatic functions. Table 27.3 lists the function of
some trace minerals.
and iron in hemoglobin). Even though they are needed in small Vitamins
amounts, animals cannot synthesize them rapidly (if at all); thus,
Normal metabolic activity depends on very small amounts of more
they must be obtained from the diet.
than a dozen organic substances called vitamins. Vitamin (L. vita,
life) is the general term for a number of chemically unrelated,
Minerals organic substances that occur in many foods in small amounts and
Some minerals are needed in relatively large amounts and are are necessary for normal metabolic functioning. Vitamins may be
called essential minerals, or macrominerals. For example, water soluble or fat soluble. Most water-soluble vitamins, such as
sodium and potassium are vital to the functioning of every the B vitamins and vitamin C, are coenzymes needed in metabolism
TA B L E 2 7 . 2 TA B L E 2 7 . 3
PHYSIOLOGICAL ROLES OF THE ESSENTIAL SOME PHYSIOLOGICAL ROLES OF TRACE MINERALS
MINERALS (MACROMINERALS) ANIMALS REQUIRE (MICROMINERALS) IN ANIMALS
IN LARGE AMOUNTS
MINERAL PHYSIOLOGICAL ROLES
MINERAL MAJOR PHYSIOLOGICAL ROLES Cobalt (Co) Component of vitamin B12; essential for red
Calcium (Ca) Component of bone and teeth; essential for blood cell production
normal blood clotting; needed for normal mus- Copper (Cu) Component of many enzymes; essential for
cle, neuron, and cellular function melanin and hemoglobin synthesis; part of
Chlorine (Cl) Principal negative ion in extracellular fluid; cytochromes
important in acid-base and fluid balance; Fluorine (F) Component of bone and teeth; prevents
needed to produce stomach HCl tooth decay
Magnesium (Mg) Component of many coenzymes; needed for Iodine (I) Component of thyroid hormones
normal neuron and muscle function, as well as
Iron (Fe) Component of hemoglobin, myoglobin,
carbohydrate and protein metabolism
enzymes, and cytochromes
Potassium (K) Major constituent of bones, blood plasma;
Manganese (Mn) Activates many enzymes; an enzyme
needed for energy metabolism
essential for urea formation and parts of
Phosphorus (P) Major positive ion in cells; influences muscle the Krebs cycle
contraction and neuron excitability; part of
Molybdenum (Mo) Constituent of some enzymes
DNA, RNA, ATP, energy metabolism
Selenium (Se) Needed in fat metabolism
Sodium (Na) Principal positive ion in extracellular fluid;
important in fluid balance; essential for con- Zinc (Zn) Component of at least 70 enzymes; needed
duction of action potentials, active transport for wound healing and fertilization
Sulfur (S) Protein structure; detoxification reactions and
other metabolic activity
in a special organ or cavity (figure 27.2b). Nutrients from the

food then pass into body cells lining the organ or cavity and can
take part in energy metabolism or biosynthesis.
(table 27.4). The fat-soluble vitamins have various functions
(table 27.5).
The dietary need for vitamin C and the fat-soluble vitamins ANIMAL STRATEGIES FOR
(A, D, E and K) tends to be limited to the vertebrates. Even in GETTING AND USING FOOD
closely related groups, vitamin requirements vary. For example,
among vertebrates, humans and guinea pigs require vitamin C, but As noted earlier, only a few protists and animals can absorb nutri-
rabbits do not. Some birds require vitamin A; others do not. ents directly from their external environment via intracellular
digestion. Most animals must work for their nutrients. The num-
ber of specializations that have evolved for food procurement
DIGESTION (feeding) and extracellular digestion are almost as numerous as
the number of animal species. What follows is a brief discussion of
In some of the simplest forms of life (the protists and sponges), the major feeding strategies animals use.
some cells take in whole food particles directly from the envi-
ronment by diffusion, active transport, and/or endocytosis and
break them down with enzymes to obtain nutrients. This strat- CONTINUOUS VERSUS
egy is called intracellular (“within the cell’’) digestion (figure DISCONTINUOUS FEEDERS
27.2a). Intracellular digestion circumvents the need for the
mechanical breakdown of food or for a gut or other cavity in One variable related to the structure of digestive systems is
which to chemically digest food. At the same time, however, whether an animal is a continuous or discontinuous feeder. Many
intracellular digestion limits an animal’s size and complexity— continuous feeders are slow-moving or completely sessile animals
only very small pieces of food can be used. Intracellular digestion (they remain permanently in one place). For example, aquatic sus-
provides all or some of the nutrients in protozoa, sponges, pension feeders, such as tube worms and barnacles, remain in one
cnidarians, platyhelminths, rotifers, bivalve molluscs, and prim- place and continuously “strain” small food particles from the water.
itive chordates. Discontinuous feeders tend to be active, sometimes highly
Larger animals have evolved structures and mechanisms mobile, animals. Typically, discontinuous feeders have more
for extracellular digestion: the enzymatic breakdown of digestive specializations than continuous feeders because discon-
larger pieces of food into constituent molecules, usually tinuous feeders take in large meals that must be either ground up
TA B L E 2 7 . 4
WATER-SOLUBLE VITAMINS
VITAMIN CHARACTERISTICS FUNCTIONS SOURCES

Thiamin (vitamin B1) Destroyed by heat and oxygen, Part of coenzyme needed for oxida- Lean meats, liver, eggs, whole-
especially in alkaline tion of carbohydrates, and coen- grain cereals, leafy green
environment zyme needed in synthesis of vegetables, legumes
ribose
Riboflavin (vitamin B2) Stable to heat, acids, and oxida- Part of enzymes and coenzymes Meats, dairy products, leafy
tion; destroyed by alkalis and needed for oxidation of glucose green vegetables, whole-
light and fatty acids and for cellular grain cereals
growth
Niacin (nicotinic acid) Stable to heat, acids, and alka- Part of coenzymes needed for oxida- Liver, lean meats, poultry,
lis; converted to niacinamide tion of glucose and synthesis of peanuts, legumes
by cells; synthesized from proteins, fats, and nucleic acids
tryptophan
Vitamin B6 Group of three compounds; sta- Coenzyme needed for synthesis of Liver, meats, fish, poultry,
ble to heat and acids; de- proteins and various amino bananas, avocados, beans,
stroyed by oxidation, alkalis, acids, for conversion of trypto- peanuts, whole-grain
and ultraviolet light phan to niacin, for production of cereals, egg yolk
antibodies, and for synthesis
of nucleic acids
Pantothenic acid Destroyed by heat, acids, and Part of coenzyme needed for oxida- Meats, fish, whole-grain cereals,
alkalis tion of carbohydrates and fats legumes, milk, fruits, vegetables
Cyanocobalamin (vitamin B12) Complex, cobalt-containing Part of coenzyme needed for synthe- Liver, meats, poultry, fish, milk,
compound; stable to heat; sis of nucleic acids and for cheese, eggs
inactivated by light, strong metabolism of carbohydrates;
acids, and strong alkalis; plays role in synthesis of myelin
absorption regulated by in-
trinsic factor from gastric
glands; stored in liver
Folate (folic acid) Occurs in several forms; de- Coenzyme needed for metabolism of Liver, leafy green vegetables,
stroyed by oxidation in acid certain amino acids and for syn- whole-grain cereals, legumes
environment or by heat in thesis of DNA; promotes produc-
alkaline environment; stored tion of normal red blood cells
in liver, where it is converted
into folinic acid
Biotin Stable to heat, acids, and light; Coenzyme needed for metabolism of Liver, egg yolk, nuts, legumes,
destroyed by oxidation and amino acids and fatty acids and mushrooms
alkalis for synthesis of nucleic acids
Ascorbic acid Closely related to monosaccha- Needed for production of collagen, Citrus fruits, citrus juices, toma-
rides; stable in acids, but conversion of folate to folinic toes, cabbage, potatoes, leafy
destroyed by oxidation, heat, acid, and metabolism of certain green vegetables, fresh fruits
light, and alkalis amino acids; promotes absorption
of iron and synthesis of hormones
from cholesterol
or stored, or both. Many carnivores, for example, pursue and cap- Herbivores spend more time eating than carnivores do, but
ture relatively large prey. When successful, they must eat large they are also discontinuous feeders. They need to move from area
meals so that they need not spend their time in the continuous to area when food is exhausted and, at least in natural environ-
pursuit of prey. Thus, carnivores have digestive systems that per- ments, must limit their grazing time to avoid excessive exposure to
mit the storage and gradual digestion of large, relatively infre- predators. Thus, their digestive systems permit relatively rapid
quent meals. food gathering and gradual digestion.
TA B L E 2 7 . 5
FAT-SOLUBLE VITAMINS
VITAMIN CHARACTERISTICS FUNCTIONS SOURCES

Vitamin A Occurs in several forms; synthesized Necessary for synthesis of visual Liver, fish, whole milk, butter, eggs, leafy
from carotenes; stored in liver; stable pigments, mucoproteins, and mu- green vegetables, and yellow and or-
in heat, acids, and alkalis; unstable copolysaccharides; for normal devel- ange vegetables and fruits
in light opment of bones and teeth; and for
maintenance of epithelial cells
Vitamin D A group of sterols; resistant to heat, oxi- Promotes absorption of calcium and Produced in skin exposed to
dation, acids, and alkalis; stored in phosphorus; promotes development ultraviolet light; in milk, egg yolk,
liver, skin, brain, spleen, and bones of teeth and bones fish-liver oils, fortified foods
Vitamin E A group of compounds; resistant to heat An antioxidant; prevents oxidation of Oils from cereal seeds, salad oils, mar-
and visible light; unstable in pres- vitamin A and polyunsaturated fatty garine, shortenings, fruits, nuts, and
ence of oxygen and ultraviolet light; acids; may help maintain stability of vegetables
stored in muscles and adipose tissue cell membranes
Vitamin K Occurs in several forms; resistant to Needed for synthesis of prothrombin; Leafy green vegetables, egg yolk, pork
heat, but destroyed by acids, alkalis, needed for blood clotting liver, soy oil, tomatoes, cauliflower
and light; stored in liver
SUSPENSION FEEDERS
Outside the
animal
Suspension feeding is the removal of suspended food particles
Nucleus
from the surrounding water by some sort of capture, trapping, or
Food particle filtration structure. This feeding strategy involves three steps:
Endocytosis (1) transport of water past the feeding structure, (2) removal of
E Nutrients nutrients from the water, and (3) transport of the nutrients to the
E
mouth of the digestive system. Sponges, ascidians, branchiopods,
Enzymes (E)
ectoprocts, entoprocts, phoronids, most bivalves, and many crus-
break down taceans, polychaetes, gastropods, and some nonvertebrate chor-
food in food dates are suspension feeders.
vacuole
(a)
DEPOSIT FEEDERS
Gut
cavity
Deposit feeding involves primarily omnivorous animals. These
animals obtain their nutrients from the sediments of soft-bottom
Nucleus
Enzymes habitats (muds and sands) or terrestrial soils. Direct deposit feed-
break down Enzymes ers simply swallow large quantities of sediment (mud, soil, sand,
food outside
the cell organic matter). The usable nutrients are digested, and the
Nutrients remains pass out the anus. Direct deposit feeding occurs in many
absorbed
polychaete annelids, some snails, some sea urchins, and in most
into cell
earthworms. Other direct deposit feeders utilize tentacle-
like structures to consume sediment. Examples include sea
cucumbers, most sipunculans, certain clams, and several types of
Nutrients polychaetes.
(b)
FIGURE 27.2 HERBIVORY

Intracellular and Extracellular Digestion. (a) A simple invertebrate,
such as a sponge, has no gut and thus carries out intracellular digestion. Tiny Herbivory (L. herba, herb ⫹ vorare, to eat) is the consumption of
food particles are taken into the body wall cells by endocytosis. Digestive macroscopic plants. This common feeding strategy requires the
enzymes in the vacuole then break the small particles into constituent mol- ability to “bite and chew’’ large pieces of plant matter (macroher-
ecules. (b) A dog, for example, has a gut and so can take in and digest (ex-
tracellularly) relatively large food particles. Cells lining the gut cavity bivory). Although biting and chewing mechanisms evolved
secrete enzymes into the cavity. There, the enzymes break down food mate- within the architectural framework of a number of invertebrate
rials into constituent nutrients, and the nearby cells absorb these nutrients. lineages, they are often characterized by the development of
hard surfaces (e.g., teeth) that powerful muscles manipulate. supplement their nutrition with organic carbon that symbiotic
Invertebrates that evolved macroherbivory include molluscs, bacteria fix within the pogonophoran’s tissues.
polychaete worms, arthropods, and sea urchins.
Many molluscs have a radula. A radula is a muscularized,
belt-like rasp armed with chitinous teeth. Molluscs use the FLUID FEEDERS
radula to scrape algae off rocks or to tear the leaves off terrestrial
The biological fluids of animals and plants are a rich source of
plants. Polychaetes have sets of large chitinous teeth on an ever-
nutrients. Feeding on this fluid is called fluid feeding. Fluid feed-
sible proboscis or pharynx that is used to scrape off algae. This
ing is especially characteristic of some parasites, such as the intes-
toothed pharynx is also suitable for carnivory when plant mate-
tinal nematodes that bite and rasp off host tissue or suck blood.
rial is scarce. Macroherbivory is found in almost every group of
External parasites (ectoparasites), such as leeches, ticks, mites,
arthropods. For example, insects and crustaceans have large,
lampreys, and certain crustaceans, use a wide variety of mouth-
powerful mandibles capable of biting off plant material and sub-
parts to feed on body fluids. For example, the sea lamprey has a
sequently grinding and chewing it before passing the plant mate-
funnel structure surrounding its mouth (see figure 27.6a). The fun-
rial to the mouth.
nel is lined with over 200 rasping teeth and a rasplike tongue. The
lamprey uses the funnel like a suction cup to grip its fish host, and
PREDATION then with its tongue, rasps a hole in the fish’s body wall. The lam-
prey then sucks blood and body fluids from the wound.
Predation (L. praedator, a plunderer, pillager) is one of the most Insects have the most highly developed sucking structures
sophisticated feeding strategies, since it requires the capture of for fluid feeding. For example, butterflies, moths, and aphids have
live prey. Only a few generalizations about the many kinds of pre- tubelike mouthparts that enable them to suck up plant fluids.
dation are presented here; discussions of various taxa are pre- Blood-sucking mosquitoes have complex mouthparts with pierc-
sented in their appropriate chapters. ing stylets.
Predators can be classified by how they capture their prey: Most pollen- and nectar-feeding birds have long bills and
motile stalkers, lurking predators, sessile opportunists, or grazers. tongues. In fact, the bill is often specialized (in shape, length, and
Motile stalkers actively pursue their prey. Examples include ciliate curvature) for particular types of flowers. The tongues of some
protozoa, nemerteans, polychaete worms, gastropods, octopuses birds have a brushlike tip or are hollow, or both, to collect the
and squids, crabs, sea stars, and many vertebrates. Lurking preda- nectar from flowers. Other nectar-feeding birds have short bills;
tors sit and wait for their prey to come within seizing distance. they make a hole in the base of a flower and use their tongue to
Examples include certain species of praying mantises, shrimp, obtain nectar through the hole.
crabs, spiders, polychaetes, and many vertebrates. Sessile oppor- The only mammals that feed exclusively on blood are the
tunists usually are not very mobile. They can only capture prey vampire bats, such as Desmodus, of tropical South and Central
when the prey organism comes into contact with them. Examples America. These bats attack birds, cattle, and horses, using knife-
include certain protozoa, barnacles, and cnidarians. Grazing car- sharp front teeth to pierce the surface blood vessels, and then lap
nivores move about the substrate picking up small organisms. at the oozing wound. Nectar-feeding bats have a long tongue to
Their diet usually consists largely of sessile and slow-moving ani- extract the nectar from flowering plants, and compared to the
mals, such as sponges, ectoprocts, tunicates, snails, worms, and blood-feeding bats, have reduced dentition. In like manner, the
small crustaceans. nectar-feeding honey possum has a long, brush-tipped tongue and
reduced dentition.
SURFACE NUTRIENT ABSORPTION

Some highly specialized animals have dispensed entirely with all DIVERSITY IN DIGESTIVE
mechanisms for prey capture, ingestion of food particles, and STRUCTURES: INVERTEBRATES
digestive processes. Instead they directly absorb nutrients from the
external medium across their body surfaces. This medium may be In primitive, multicellular animals, such as cnidarians, the gut is a
nutrient-rich seawater, fluid in other animals’ digestive tracts, or blind (closed) sac called a gastrovascular cavity. It has only one
the body fluids of other animals. For example, some free-living opening that is both entrance and exit (figure 27.3a); thus, it is an
protozoa, such as Chilomonas, absorb all of their nutrients across incomplete digestive tract. Some specialized cells in the cavity
their body surface. The endoparasitic protozoa, cestode worms, secrete digestive enzymes that begin the process of extracellular
endoparasitic gastropods, and crustaceans (all of which lack digestion. Other phagocytic cells that line the cavity engulf food
mouths and digestive systems) also absorb all of their nutrients material and continue intracellular digestion inside food vacuoles.
across their body surface. Some flatworms have similar digestive patterns (figure 27.3b).
A few nonparasitic multicellular animals also lack a mouth The development of the anus and complete digestive tract
and digestive system and absorb nutrients across their body in the aschelminths was an evolutionary breakthrough (figure
surface. Examples include the gutless bivalves and pogonoph- 27.3c). A complete digestive tract permits the one-way flow of
oran worms. Interestingly, many pogonophoran worms absorb ingested food without mixing it with previously ingested food or
some nutrients from seawater across their body surface and also waste. Complete digestive tracts also have the advantage of
Mouth Wastes
Anus Pharynx
Food Eyespot
Tentacles
with
nematocysts Mouth
Gastrovascular
cavity
Gastrovascular (c) Nematode

cavity
Crop Stomach
Mouth
Gland Rectum
Pharynx
cell Esophagus
Pseudopod Mouth
Anus
Food
particle Opening of
pharynx
Gastric Intestine
Food caeca
vacuole
(a) Cnidarian (Hydra) (b) Planarian (d) Insect
FIGURE 27.3
Various Types of Digestive Structures in Invertebrates. (a) The gastrovascular cavity of cnidarian (Hydra) is an incomplete digestive tract because
its one opening, a mouth, must serve as the entry and exit point for food and waste. Extracellular digestion occurs in the gastrovascular cavity, and
intracellular digestion occurs inside food vacuoles formed when phagocytic cells engulf food particles. (b) Even though the gastrovascular cavity in a
platyhelminth (planarian) branches extensively, it is also an incomplete digestive tract with only one opening. When a planarian feeds, it sticks its
muscular pharynx out of its mouth and sucks in food. (c) A nematode (Ascaris) has a complete digestive tract with a mouth, pharynx, and anus. (d) The
complete digestive tract of an insect (grasshopper) has an expanded region called a crop that functions as a food storage organ.
progressive digestive processing in specialized regions along

Food
the system. Food can be digested efficiently in a series of dis-
tinctly different steps. The many variations of the basic plan of a Buccal cavity
complete digestive tract correlate with different food-gathering Cytostome

mechanisms and diets (figure 27.3d). Most of these have been pre- 1. Food vacuole
sented in the discussion of the many different protists and inver- forming at
tebrates and are not repeated in this chapter. Instead, three exam- cytopharynx
ples further illustrate digestive systems in protozoa and 2. Excess water

invertebrates: (1) the incomplete digestive system of a ciliated leaves food
vacuole
protozoan is an example of an intracellular digestive system; (2)
pH acid
the bivalve mollusc is an example of an invertebrate that has both 3. Lysosomes
delivering
intracellular and extracellular digestion; and (3) an insect is an enzymes to
example of an invertebrate that has extracellular digestion and a food vacuole pH alkaline
complete digestive tract. 4. Food particles
undergoing pH alkaline
digestion in
PROTOZOA vacuole
5. Residual
As presented in chapter 8, protozoa may be autotrophic, saprozoic, vacuole
or heterotrophic (ingest food particles). Ciliated protozoa are Cytopyge
good examples of protists that utilize heterotrophic nutrition. Cil-
Waste from
iary action directs food from the environment into the buccal vacuole
cavity and cytostome (figure 27.4). The cytostome opens into
the cytopharynx, which enlarges as food enters and pinches off a
food-containing vacuole. The detached food vacuole then moves FIGURE 27.4
through the cytoplasm. During this movement, excess water is Intracellular Digestion in a Ciliated Protozoan. Cilia direct food to-
ward the cytostome (“mouth’’). The food enters the cytopharynx, where
removed from the vacuole, the contents are acidified and then a food vacuole forms and detaches from the cytopharynx. The detached
made alkaline and a lysosome adds digestive enzymes. The food vacuole undergoes acidic and alkaline digestion, and the waste vacuole
particles are then digested within the vacuole and the nutrients moves to the cytopyge (“anus’’) for excretion.
(a)
Food
(b)
Fragmentation
spherules
Large food
particles
Digestive
diverticula Right pouch
Major
Left typhlosole
pouch
Midgut
Style sac
Crystalline
(e) style (c)
Esophagus
Gastric shield
(d)
FIGURE 27.5
Extracellular and Intracellular Digestion in a Bivalve Mollusc. (a) Extracellular digestion begins before food ingestion by the dissolving of the crys-
talline style and the formation of fragmentation spherules in the stomach. (b) As food enters the stomach, the rotating style and the enzymes released
by the gastric shield mechanically and enzymatically break it down. (c) The small food particles then move into the digestive diverticulae for intracel-
lular digestion. (d) A progressive passage of food particles from the stomach to the digestive diverticulae follows cessation of feeding. (e) During this
resting phase, the stomach empties and the style reforms, while intracellular digestion in the diverticulae is completed, and fragmentation spherules be-
gin to form again. The movement of fragmentation spherules starts the next feeding cycle.
absorbed into the cytoplasm. The residual vacuole then excretes and enzymatic breakdown of food during feeding provides the
its waste products via the cytopyge. small particles for intracellular digestion. Intracellular digestion
releases the nutrients into the blood and produces the fragmenta-
tion spherules that both excrete wastes and lower the pH for opti-
BIVALVE MOLLUSCS mal extracellular digestion. These three cycles are linked to tidal
immersion and emersion of the mollusc.
Many bivalve molluscs suspension feed and ingest small food parti-
cles. The digestive tract has a short esophagus opening into a stom-
ach, midgut, hindgut, and rectum. The stomach contains a crys- INSECTS
talline style, gastric shield, and diverticulated region. These
diverticulae are blind-ending sacs that increase the surface area for The grasshopper is a representative insect with a complete diges-
absorption and intracellular digestion. The midgut, hindgut, and rec- tive tract and extracellular digestion (see figure 27.3d). During
tum function in extracellular digestion and absorption (figure 27.5). feeding, the mandibles and maxillae first break up (masticate) the
Digestion is a coordination of three cycles: (1) feeding, (2) food, which is then taken into mouth and passed to the crop via
extracellular digestion, and (3) intracellular digestion. The resting the esophagus. During mastication, the salivary glands add saliva
phase is preparative for extracellular digestion. The mechanical to the food to lubricate it for passage through the digestive tract.
Saliva also contains the enzyme amylase, which begins the enzy- whether an animal feeds on plants or animals, and on how it
matic digestion of carbohydrates. This digestion continues during obtains its food. The teeth of snakes slope backward to aid
food storage in the crop. The midgut secretes other enzymes (car- in the retention of prey while swallowing (figure 27.7a), and
bohydrases, lipases, proteases) that enter the crop. Food passes the canine teeth of wolves are specialized for ripping food
slowly from the crop to the stomach, where it is mechanically (figure 27.7b). Herbivores, such as deer, have predominantly
reduced and the nutrient particles sorted. Large particles are grinding teeth, the front teeth of a beaver are used for chiseling
returned to the crop for further processing; the small particles trees and branches, and the elephant has two of its upper, front
enter the gastric cecae, where extracellular digestion is completed. teeth specialized as weapons and for moving objects (figure
Most nutrient absorption then occurs in the intestine. Undigested 27.7c–e). Because humans, pigs, bears, raccoons, and a few
food is moved along the intestine and passes into the rectum, other mammals are omnivores, they have teeth that can per-
where water and ions are absorbed. The solid fecal pellets that form a number of tasks—tearing, ripping, chiseling, and grind-
form then pass out of the animal via the anus. During this entire ing (figure 27.7f).
feeding process the nervous system, the endocrine system, and the
presence of food exert considerable control over enzyme produc-
tion at various points in the digestive tract. SALIVARY GLANDS
Most fishes lack salivary glands in the head region. Lampreys are
DIVERSITY IN DIGESTIVE an exception because they have a pair of glands that secrete an
anticoagulant needed to keep their prey’s blood flowing as they
STRUCTURES: VERTEBRATES feed. Modified salivary glands of some snakes produce venom that
The complete vertebrate digestive tract (gut tube) is highly spe- is injected through fangs to immobilize prey. Because the secretion
cialized in both structure and function for the digestion of a wide of oral digestive enzymes is not an important function in amphib-
variety of foods. The basic structures of the gut tube include the ians or reptiles, salivary glands are absent. Most birds lack salivary
buccal cavity, pharynx, esophagus, stomach, small intestine, large glands, while all mammals have them.
intestine, rectum, and anus/cloaca. In addition, three important
glandular systems are associated with the digestive tract: (1) the
salivary glands; (2) the liver, gallbladder, and bile duct; and (3) ESOPHAGI
the pancreas and pancreatic duct. The esophagus (pl., esophagi) is short in fishes and amphibians,
Because most vertebrates spend the majority of their time but much longer in amniotes due to their longer necks. Grain-
acquiring food, feeding is the universal pastime. The oral cavity and seed-eating birds have a crop that develops from the caudal
(mouth), teeth, intestines, and other major digestive structures portion of the esophagus (figure 27.8a). Storing food in the crop
usually reflect the way an animal gathers food, the type of food it ensures an almost continuous supply of food to the stomach and
eats, and the way it digests that food. These major digestive struc- intestine for digestion. This structure allows these birds to
tures are now discussed to illustrate the diversity of form and func- reduce the frequency of feeding and still maintain a high meta-
tion among different vertebrates. bolic rate.
TONGUES
STOMACHS
A tongue or tonguelike structure develops in the floor of the oral
cavity in many vertebrates. For example, a lamprey has a protrusi- The stomach is an ancestral vertebrate structure that evolved as
ble tongue with horny teeth that rasp its prey’s flesh (figure 27.6a). vertebrates began to feed on larger organisms that were
Fishes may have a primary tongue that bears teeth that help hold caught at less frequent intervals and required storage. Some
prey; however, this type of tongue is not muscular (figure 27.6b). zoologists believe that the gastric glands and their production
Tetrapods have evolved mobile tongues for gathering food. Frogs of hydrochloric acid (HCl) evolved in the context of killing bac-
and salamanders and some lizards can rapidly project part of their teria and helping preserve food. The enzyme pepsinogen may
tongue from the mouth to capture an insect (figure 27.6c). A have evolved later because the stomach is not essential for
woodpecker has a long, spiny tongue for gathering insects and digestion.
grubs (figure 27.6d). Ant- and termite-eating mammals also gather
food with long, sticky tongues. Spiny papillae on the tongues of GIZZARDS
cats and other carnivores help these animals rasp flesh from a bone.
Some fishes, some reptiles such as crocodilians, and all birds have
TEETH a gizzard for grinding up food (figure 27.8a). The bird’s gizzard
develops from the posterior part of the stomach called the ven-
With the exception of birds, turtles, and baleen whales, most triculus. Pebbles (grit) that have been swallowed are often
vertebrates have teeth. Birds lack teeth, probably to reduce retained in the gizzard of grain-eating birds and facilitate the
body weight for flight. Teeth are specialized, depending on grinding process.
(b)
(a)
(c)
FIGURE 27.6
Tongues. (a) Rasping tongue and mouth of a lamprey. (b) Fish tongue.
(c) Tongue of a chameleon catching an insect. (d) The tongue of a
woodpecker extracts insects from the bark of a tree. (d)
RUMENS In ruminants, the upper portion of the stomach expands to

form a large pouch, the rumen, and a smaller reticulum. The lower
Ruminant mammals—animals that “chew their cud,’’ such as portion of the stomach consists of a small antechamber, the oma-
cows, sheep, and deer—show some of the most unusual modifi- sum, with a “true’’ stomach, or abomasum, behind it (figure 27.9).
cations of the stomach. This method of digestion has evolved Food first enters the rumen, where it encounters the microorgan-
in animals that need to eat large amounts of food relatively isms. Aided by copious fluid secretions, body heat, and churning of
quickly, but can chew the food at a more comfortable or safer the rumen, the microorganisms partially digest the food and reduce
location. More important though, the ruminant stomach pro- it to a pulpy mass. Later, the pulpy mass moves into the reticulum,
vides an opportunity for large numbers of microorganisms to from which mouthfuls are regurgitated as “cud’’ (L. ruminare, to
digest the cellulose walls of grass and other vegetation. Cellu- chew the cud). At this time, food is thoroughly chewed for the
lose contains a large amount of energy; however, animals gener- first time. When reswallowed, the food enters the rumen, where it
ally lack the ability to produce the enzyme cellulase for digesting becomes more liquid in consistency. When it is very liquid, the
cellulose and obtaining its energy. Because gut microorganisms digested food material flows out of the reticulum and into the
can produce cellulase, they have made the herbivorous lifestyle omasum and then the glandular region, the abomasum. Here the
more effective. digestive enzymes are first encountered, and digestion continues.
Esophagus
Crop
(a)
Stomach
Liver
Gizzard
Pancreas
(b)
Small
intestine
Rectum
Cloaca
(a)
(c)
Pharynx Stomach Pyloric valve Anus
Lung duct Bile duct Spiral valve Intestine
(d) (b)
Duct to swim bladder Gallbladder Stomach
Pharynx Pyloric ceca Intestine Anus
(c)
(e) Pharynx
Liver Esophagus
Stomach
Gallbladder
Small
intestine
Duodenum
Pylorus
Bladder
Cloaca
(f) (d)
FIGURE 27.8
Arrangement of Stomachs and Intestines in a Variety of Vertebrates.
FIGURE 27.7 (a) Pigeon. (b) Lungfish. (c) Teleost fish. (d) Frog.
Arrangement of Teeth in a Variety of Vertebrates. (a) Snake.
(b) Wolf. (c) Deer. (d) Beaver. (e) Elephant. (f) Human.
Ascending
colon
Ileum
Cecum
Rumen Esophagus Omasum Appendix
FIGURE 27.10
Extensive Cecum of a Nonruminant Herbivore, Such as a Rabbit.
Pylorus
The cecum contains microorganisms that produce digestive enzymes
(e.g., cellulase that helps break down cellulose).
ments that enter the circulation. These pigments are subsequently

extracted from the circulation in the liver and excreted in the bile
Reticulum
as bilirubin (“red bile’’) and biliverdin (“green bile’’).
Because of the importance of bile in fat digestion, the gall-
Abomasum
(true stomach) bladder is relatively large in carnivores and vertebrates, in which
fat is an important part of the diet. It is much reduced or absent in
FIGURE 27.9 bloodsuckers, such as the lamprey, and in animals that feed pri-
Ruminant Mammal. Four-chambered stomach of a cow, where symbi- marily on plant food (e.g., some teleosts, many birds, and rats).
otic microorganisms digest cellulose.
PANCREATA
CECA Every vertebrate has a pancreas (pl., pancreata); however, in lam-
preys and lungfishes it is embedded in the wall of the intestine and is
Microorganisms attack the food of ruminants before gastric diges-
not a visible organ. Both endocrine and exocrine tissues are present,
tion, but in the typical nonruminant herbivore, microbial action
but the cell composition varies. Pancreatic fluid containing many
on cellulose occurs after digestion. Rabbits, horses, and rats digest
enzymes empties into the small intestine via the pancreatic duct.
cellulose by maintaining a population of microorganisms in their
unusually large cecum, the blind pouch that extends from the
colon (figure 27.10). Adding to this efficiency, a few non-ruminant
herbivores, such as mice and rabbits, eat some of their own feces INTESTINES
to process the remaining materials in them, such as vitamins.
The configuration and divisions of the small and large intestines
vary greatly among vertebrates. Intestines are closely related to
LIVERS AND GALLBLADDERS the animal’s type of food, body size, and levels of activity. For
example, cyclostomes, chondrichthian fishes, and primitive bony
In those vertebrates with a gallbladder, it is closely associated with fishes have short, nearly straight intestines that extend from the
the liver. The liver manufactures bile, which the gallbladder then stomach to the anus (see figure 27.8b). In more advanced bony
stores. Bile is a fluid containing bile salts and bile pigments. Bile fishes, the intestine increases in length and begins to coil (see
salts play an important role in the digestion of fats, although they figure 27.8c). The intestines are moderately long in most amphib-
are not digestive enzymes. They emulsify dietary fat, breaking it ians and reptiles (see figure 27.8d) In birds and mammals, the
into small globules (emulsification) on the surface of which the intestines are longer and have more surface area than those of
fat-digesting enzyme lipase can function. Bile pigments result from other tetrapods (see figure 27.8a). Birds typically have two ceca,
phagocytosis of red blood cells in the spleen, liver, and red bone and mammals have a single cecum at the beginning of the large
marrow. Phagocytosis cleaves the hemoglobin molecule, releasing intestine. The large intestine is much longer in mammals than in
iron, and the remainder of the molecule is converted into pig- birds, and it empties into the cloaca in most vertebrates.
Muscle layer
Serosa Longitudinal Circular Mucosa

smooth muscle smooth muscle
Submucosa Lumen
FIGURE 27.12
Mammalian Gastrointestinal Tract. Common structural layers of the
gastrointestinal tract. The central lumen extends from the mouth to
the anus.
The process of digesting and absorbing nutrients in a mam-

mal includes:
1. Ingestion—eating
2. Peristalsis—the involuntary, sequential muscular contrac-
tions that move ingested nutrients along the digestive
tract
3. Segmentation—mixing the contents in the digestive tract
4. Secretion—the release of hormones, enzymes, and specific
ions and chemicals that take part in digestion
5. Digestion—the conversion of large nutrient particles or
molecules into small particles or molecules
6. Absorption—the passage of usable nutrient molecules from
the small intestine into the bloodstream and lymphatic sys-
tem for the final passage to body cells
7. Defecation—the elimination from the body of undigested
and unabsorbed material as waste
FIGURE 27.11
Major Organs and Parts of the Human Digestive System. Food GASTROINTESTINAL MOTILITY
passes from the mouth through the pharynx and esophagus to the stom- AND ITS CONTROL
ach. From the stomach, it passes to the small intestine, where nutrients
are broken down and absorbed into the circulatory and lymphatic sys- As with any organ, the function of the gastrointestinal tract is
tems. Nutrients then move to the large intestine, where water is reab- determined by the type of tissues it contains. Most of the mam-
sorbed, and feces form. Feces exit the body via the anal canal.
malian gastrointestinal tract has the same anatomical structure
along its entire length (figure 27.12). From the outside inward is a
thin layer of connective tissue called the serosa. (The serosa forms
a moist epithelial sheet called the peritoneum. This peritoneum
lines the entire abdominal cavity and covers all internal organs.
THE MAMMALIAN DIGESTIVE The space it encompasses is the coelom.) Next are the longitudi-
SYSTEM nal smooth-muscle layer and circular smooth-muscle layer.
Underneath this muscle layer is the submucosa. The submucosa
Humans, pigs, bears, raccoons, and a few other mammals are contains connective tissue, blood, and lymphatic vessels. The
omnivores. The digestive system of an omnivore has the mechan- mucosa faces the central opening, which is called a lumen.
ical and chemical ability to process many kinds of foods. The sec- The coordinated contractions of the muscle layers of the
tions that follow examine the control of gastrointestinal motility, gastrointestinal tract mix the food material with various secre-
the major parts of the alimentary canal, and the accessory organs tions and move the food from the oral cavity to the rectum. The
of digestion (figure 27.11). two types of movement involved are peristalsis and segmentation.
FIGURE 27.13
Peristalsis and Segmentation. (a) Peristaltic waves move food through the esophagus to the stomach. (b) In segmentation, simultaneous muscular
contractions of many sections of the intestine (blue arrows) help mix nutrients with digestive secretions.
During peristalsis (Gr. peri, around ⫹ stalsis, contraction), smooth or skeletal muscle at the beginning or ends of specific
food advances through the gastrointestinal tract when the rings of regions of the gut tract. For example, the cardiac sphincter is
circular smooth muscle contract behind it and relax in front of it between the esophagus and stomach, and the pyloric sphincter is
(figure 27.13a). Peristalsis is analogous to squeezing icing from a between the stomach and small intestine.
pastry tube. The small and large intestines also have rings of Control of gastrointestinal activity is based on the volume
smooth muscles that repeatedly contract and relax, creating an and composition of food in the lumen of the gut. For example,
oscillating back-and-forth movement in the same place, called ingested food distends the gut and stimulates mechanical recep-
segmentation (figure 27.13b). This movement mixes the food with tors in the gut wall. In addition, digestion of carbohydrates, lipids,
digestive secretions and increases the efficiency of absorption. and proteins stimulates various chemical receptors in the gut wall.
Sphincters also influence the flow of material through the Signals from these mechanical and chemical stimuli travel
gastrointestinal tract and prevent backflow. Sphincters are rings of through nerve plexuses in the gut wall to control the muscular
(a)
FIGURE 27.14
Teeth. (a) The teeth of a carnivore, such as this wolf, are specialized
for slicing, puncturing, tearing, and grinding animal flesh. (b) Anatomy
of a typical mammalian tooth.
contraction that leads to peristalsis and segmentation, as well as

the secretion of various substances (e.g., mucus, enzymes) into the
gut lumen. In addition to this local control, long-distance nerve
(b)
pathways connect the receptors and effectors with the central
nervous system. Either or both of these pathways function to
maintain homeostasis in the gut. The endocrine cells of the gas-
trointestinal tract also produce hormones that help regulate secre-
tion, digestion, and absorption.
porarily seals off the opening (glottis) to the trachea so that swal-
lowed food does not enter the trachea. Initiation of the swallow-
ORAL CAVITY ing reflex can be voluntary, but most of the time it is involuntary.
When swallowing begins, sequential, involuntary contractions of
A pair of lips protects the oral cavity (mouth). The lips are highly smooth muscles in the walls of the esophagus propel the bolus or
vascularized, skeletal muscle tissue with an abundance of sensory liquid to the stomach. Neither the pharynx nor the esophagus
nerve endings. Lips help retain food as it is being chewed and play contribute to digestion.
a role in phonation (the modification of sound).
The oral cavity contains the tongue and teeth (figure
27.14). Mammals can mechanically process a wide range of foods STOMACH
because their teeth are covered with enamel (the hardest material
The mammalian stomach is a muscular, distensible sac with three
in the body) and because their jaws and teeth exert a strong force.
main functions. It (1) stores and mixes the food bolus received
The oral cavity is continuously bathed by saliva, a watery fluid
from the esophagus, (2) secretes substances (enzymes, mucus, and
that at least three pairs of salivary glands secrete. Saliva moistens
hydrochloric acid [HCl]) that start the digestion of proteins, and
food, binds it with mucins (glycoproteins), and forms the ingested
(3) helps control the rate at which food moves into the small
food into a moist mass called a bolus. Saliva also contains bicar-
intestine via the pyloric sphincter (figure 27.15a).
bonate ions (HCO3⫺), which buffer chemicals in the mouth, and
The stomach is made up of an inner mucous membrane con-
thiocyanate ions (SCN⫺) and the enzyme lysozyme, which kill
taining thousands of gastric glands (figure 27.15b). Three types of
microorganisms. It also contributes an enzyme (amylase) neces-
cells are in these glands. Parietal cells secrete a solution containing
sary for the initiation of carbohydrate digestion.
HCl, and chief cells secrete pepsinogen, the precursor of the
enzyme pepsin. Both of the cells are in the pits of the gastric glands.
PHARYNX AND ESOPHAGUS The surface of the mucous membrane at the openings of the glands
contains numerous mucous cells that secrete mucus that coats the
Chapter 26 discusses how both air and swallowed foods and liquids surface of the stomach and protects it from the HCl and digestive
pass from the mouth into the pharynx—the common passageway enzymes. The surfaces of the upper gastrointestinal tract—the
for both the digestive and respiratory tracts. The epiglottis tem- esophagus and mouth—have a much thinner mucous-cell layer
Esophagus Fundic region

of stomach
Cardiac region
of stomach
Pyloric sphincter
(pylorus)
Body of
stomach
Duodenum
Pyloric canal
Pyloric region
of stomach
(a)
Gastric pits
Mucous cell
Mucous membrane
Gastric gland
Mucosal
muscles
Parietal cell
Chief cell Submucosa
Loechel
(b)
FIGURE 27.15
Stomach. (a) Food enters the stomach from the esophagus. (b) Gastric glands cover the mucosa of the stomach and include mucous cells, parietal
cells, and chief cells. Each type produces a different secretion.
than the stomach, which is why vomiting can cause a burning sen- the villi (figure 27.16c). From the lacteals, the chylomicrons move
sation in the esophagus or mouth. Endocrine cells in one part of into the lymphatics and eventually into the bloodstream for trans-
the stomach mucosa release the hormone gastrin, which travels to port throughout the body.
target cells in the gastric glands, further stimulating them. Besides absorbing organic molecules, the small intestine ab-
When the bolus of food enters the stomach, it distends the sorbs water and dissolved mineral ions. The small intestine
walls of the stomach. This distention, as well as the act of eating, absorbs about 9 liters of water per day, and the large intestine ab-
causes the gastric pits to secrete HCl (as H⫹ and Cl⫺) and sorbs the rest.
pepsinogen. The H⫹ ions cause pepsinogen to be converted into
the active enzyme pepsin. As pepsin, mucus, and HCl mix with
and begin to break down proteins, smooth mucosal muscles con- LARGE INTESTINE
tract and vigorously churn and mix the food bolus. About three to
four hours after a meal, the stomach contents have been suffi- Unlike the small intestine, the large intestine has no circular
ciently mixed and are a semiliquid mass called chyme (Gr. chy- folds, villi, or microvilli; thus, the surface area is much smaller.
mos, juice). The pyloric sphincter regulates the release of the The small intestine joins the large intestine near a blind-ended
chyme into the small intestine. sac, the cecum (L. caecum, blind gut) (see figure 27.11). The
When the stomach is empty, peristaltic waves cease; how- human cecum and its extension, the appendix, are storage
ever, after about 10 hours of fasting, new waves may occur in the sites and possibly represent evolutionary remains of a larger,
upper region of the stomach. These waves can cause “hunger functional cecum, such as is found in herbivores (see figure
pangs’’ as sensory nerve fibers carry impulses to the brain. 27.10). The appendix contains an abundance of lymphoid tissue
and may function as part of the immune system.
The major functions of the large intestine include the
SMALL INTESTINE: MAIN SITE reabsorption of water and minerals, and the formation and
OF DIGESTION storage of feces. As peristaltic waves move food residue along,
minerals diffuse or are actively transported from the residue
Most of the food a mammal ingests is digested and absorbed in the across the epithelial surface of the large intestine into the
small intestine. The human small intestine is about 4 cm in diam- bloodstream. Water follows osmotically and returns to the lym-
eter and 7 to 8 m in length (see figure 27.11). It is intermediate in phatic system and bloodstream. When water reabsorption is
length between the small intestines of typical carnivores and her- insufficient, diarrhea (Gr. rhein, to flow) results. If too much
bivores of similar size, and it reflects the human’s omnivorous eat- water is reabsorbed, fecal matter becomes too thick, resulting in
ing habits. The length of the small intestine directly relates to the constipation.
total surface area available for absorbing nutrients, as determined Many bacteria and fungi exist symbiotically in the large
by the many circular folds and minute projections of the inner gut intestine. They feed on the food residue and further break down
surface (figure 27.16a). On the circular folds, thousands of finger- its organic molecules to waste products. In turn, they secrete
like projections called villi (L. villus, tuft of hair) (sing. villus) amino acids and vitamin K, which the host’s gut absorbs. What
project from each square centimeter of mucosa (figure 27.16b,c). remains—feces—is a mixture of bacteria, fungi, undigested plant
Simple columnar epithelial cells, each bearing numerous fiber, sloughed-off intestinal cells, and other waste products.
microvilli, cover both the circular folds and villi (figure 27.16d).
These minute projections are so dense that the inner wall of the
human small intestine has a total surface area of approximately ROLE OF THE PANCREAS
300 m2—the size of a tennis court. IN DIGESTION
The first part of the small intestine, called the duodenum,
functions primarily in digestion. The next part is the jejunum, and The pancreas (Gr. pan, all ⫹ kreas, flesh) is an organ that lies
the last part is the ileum. Both function in nutrient absorption. just ventral to the stomach and has both endocrine and exocrine
The duodenum contains many digestive enzymes that intes- functions. Exocrine cells in the pancreas secrete digestive
tinal glands in the duodenal mucosa secrete. The pancreas enzymes into the pancreatic duct, which merges with the
secretes other enzymes. In the duodenum, digestion of carbohy- hepatic duct from the liver to form a common bile duct that
drates and proteins is completed, and most lipids are digested. The enters the duodenum. Pancreatic enzymes complete the diges-
jejunum and ileum absorb the end products of digestion (amino tion of carbohydrates and proteins and initiate the digestion of
acids, simple sugars, fatty acids, glycerol, nucleotides, water). lipids. Trypsin, carboxypeptidase, and chymotrypsin digest pro-
Much of this absorption involves active transport and the sodium- teins into small peptides and individual amino acids. Pancreatic
dependent ATPase pump. Sugars and amino acids are absorbed lipases split triglycerides into smaller, absorbable glycerol and
into the capillaries of the villi, whereas free fatty acids enter the free fatty acids. Pancreatic amylase converts polysaccharides
epithelial cells of the villi and recombine with glycerol to form into disaccharides and monosaccharides. Table 27.6 summarizes
triglycerides. The triglycerides are coated with proteins to form the major glands, secretions, and enzymes of the mammalian
small droplets called chylomicrons, which enter the lacteals of digestive system.
(a)
(c)
(b)
FIGURE 27.16
Small Intestine. The small intestine absorbs food over a large surface
area. (a) The lining of the intestine has many circular folds. (b,d) Fin-
gerlike villi line the intestine. A single villus contains a central capillary
network and a lymphatic lacteal, both of which transport nutrients ab-
sorbed from the lumen of the intestine. (c) The plasma membrane of the
simple columnar epithelial cells covering the villi fold into microvilli
(arrow), which further increase the surface area facing the lumen. (d)
TA B L E 2 7 . 6
MAJOR DIGESTIVE GLANDS, SECRETIONS, AND ENZYMES IN MAMMALS
PLACE OF DIGESTION SOURCE SECRETION ENZYME DIGESTIVE FUNCTION

Mouth Salivary glands Saliva Salivary amylase Begins the digestion of carbohydrates; inactivated by
stomach HCl
Mucous glands Mucus — Lubricates food bolus
Esophagus Mucous glands Mucus — Lubricates food bolus
Stomach Gastric glands Gastric juice Lipase Digests lipids into fatty acids and glycerol
Pepsin Digests proteins into polypeptides
Gastric mucosa HCl — Converts pepsinogen into active pepsin; kills
microorganisms
Mucous glands Mucus — Lubricates
Small intestine Liver Bile — Emulsifies lipids; activates lipase
Pancreas Pancreatic juice Amylase Digests starch into maltose
Chymotrypsin Digests proteins into peptides and amino acids
Lipase Digests lipids into fatty acids and glycerol (requires
bile salts)
Nuclease Digests nucleic acids into mononucleotides
Trypsin Digests proteins into peptides and amino acids
Intestinal glands Intestinal juice Enterokinase Digests inactive trypsinogen into active trypsin
Lactase Digests lactose into glucose and galactose
Maltase Digests maltose into glucose
Peptidase Digests polypeptides into amino acids
Sucrase Digests sucrose into glucose and fructose
Mucous glands Mucus — Lubricates
Large intestine Mucous glands Mucus — Lubricates
The pancreas also secretes bicarbonate (HCO3⫺) ions that 3. Manufacture of most of the plasma proteins, formation of
help neutralize the acidic food residue coming from the stomach. fetal erythrocytes, destruction of worn-out erythrocytes, and
Bicarbonate raises the pH from 2 to 7 for optimal digestion. synthesis of the blood-clotting agents prothrombin and fi-
Without such neutralization, pancreatic enzymes could not brinogen from amino acids.
function. 4. Synthesis of nonessential amino acids.
5. Conversion of galactose and fructose to glucose.
6. Oxidation of fatty acids.
7. Formation of lipoproteins, cholesterol, and phospholipids
ROLE OF THE LIVER AND (essential cell membrane components).
GALLBLADDER IN DIGESTION 8. Conversion of carbohydrates and proteins into fat.
9. Modification of waste products, toxic drugs, and poisons
The liver, the largest organ in the mammalian body, is just under
(detoxification).
the diaphragm (see figure 27.11). In the liver, millions of special-
10. Synthesis of vitamin A from carotene, and with the kidneys,
ized cells called hepatocytes take up nutrients absorbed from the
participation in the activation of vitamin D.
intestines and release them into the bloodstream. Hepatocytes
11. Maintenance of a stable body temperature by raising the
also manufacture the blood proteins prothrombin and albumin.
temperature of the blood passing through it. Its many meta-
In addition, some major metabolic functions of the liver
bolic activities make the liver the major heat producer in a
include:
mammal’s body.
1. Removal of amino acids from organic compounds. 12. Manufacture of bile salts, which are used in the small intes-
2. Urea formation from proteins and conversion of excess tine for the emulsification and absorption of simple fats,
amino acids into urea to decrease body levels of ammonia. cholesterol, phospholipids, and lipoproteins.
13. Main storage center. The liver stores glucose in the form of greenish fluid called bile that the liver cells continuously produce.
glycogen, and with the help of insulin and enzymes, converts Bile is very alkaline and contains pigments, cholesterol, lecithin,
glycogen back into glucose as the body needs it. The liver mucin, bilirubin, and bile salts that act as detergents to emulsify
also stores fat-soluble vitamins (A, D, E, and K), and miner- fats (form them into droplets suspended in water) and aid in fat
als, such as iron, from the diet. The liver can also store fats digestion and absorption. (Recall that fats are insoluble in water.)
and amino acids, and convert them into usable glucose as Bile salts also combine with the end products of fat digestion to
required. form micelles. Micelles are lipid aggregates (fatty acids and glyc-
erol) with a surface coat of bile salts. Because they are so small,
The gallbladder (L. galbinus, greenish yellow) is a small
they can cross the microvilli of the intestinal epithelium.
organ near the liver (see figure 27.11). The gallbladder stores the
SUMMARY makes fats soluble. The liver has many diverse functions. It controls
the fate of newly synthesized food molecules, stores excess glucose as
1. Nutrition describes all of those processes by which an animal takes glycogen, synthesizes many blood proteins, and converts nitrogenous
in, digests, absorbs, stores, and uses food (nutrients) to meet its and other wastes into a form that the kidneys can excrete.
metabolic needs. Digestion is the mechanical and chemical break- 9. The large intestine has little digestive or nutrient absorptive activ-
down of food into smaller particles that the individual cells of an ity. It functions principally to absorb water, to compact the mate-
animal can absorb. rial left over from digestion, and to be a storehouse for
2. Losses of biosynthetic abilities have marked much of animal evolu- microorganisms.
tion. This tendency has led to the evolution of the following nutri-
tional types: insectivores, herbivores, carnivores, and omnivores.
3. The nutrients that a heterotroph ingests can be divided into SELECTED KEY TERMS
macronutrients and micronutrients. Macronutrients are needed in
large quantities and include the carbohydrates, lipids, and proteins. autotrophs (p. 434) micronutrients (p. 434)
Micronutrients are needed in small quantities and include the vita- carnivores (p. 434) nutrition (p. 433)
mins and minerals. digestion (p. 433) omnivores (p. 434)
herbivores (p. 434) suspension feeding (p. 438)
4. Only a few protists and animals can absorb nutrients directly from
heterotrophs (p. 434) vitamin (p. 435)
their external environment. Most animals must work for their nu-
macronutrients (p. 434)
trients. Various specializations have evolved for food procurement
(feeding) in animals. Some examples include continuous versus dis-
continuous feeding, suspension feeding, deposit feeding, herbivory,
predation, surface nutrient absorption, and fluid feeding. CRITICAL THINKING QUESTIONS
5. The evolution and structure of the digestive system in various in- 1. What advantages are there to digestion? Would it not be simpler for
vertebrates and vertebrates reflect their eating habits, their rate of a vertebrate to simply absorb carbohydrates, lipids, and proteins from
metabolism, and their body size. its food and to use these molecules without breaking them down?
6. The digestive system of vertebrates is one-way, leading from the 2. What might have been some evolutionary pressures acting on ani-
mouth (oral cavity), to the pharynx, esophagus, stomach, small in- mals that led to the internalization of digestive systems?
testine, large intestine, rectum, and anus. 3. Many digestive enzymes are produced in the pancreas and released
7. The coordinated contractions of the muscle layer of the gastroin- into the duodenum. Why, then, has the mammalian stomach
testinal tract mix food material with various secretions and move evolved the ability to produce pepsinogen?
the food from the oral cavity to the rectum. The two types of 4. Human vegetarians, unlike true herbivores, have no highly special-
movement involved are segmentation and peristalsis. ized fermentation chambers. Why?
8. Most digestion occurs in the duodenal portion of the small intestine. 5. Trace the fate of a hamburger from the mouth to the anus, identify-
The products of digestion are absorbed in the walls of the jejunum ing sites and mechanisms of digestion and absorption.
and ileum. In the process of digestion, bile that the liver secretes
ONLINE LEARNING CENTER • RELATED WEB LINKS

Digestion and Nutrition
Visit our Online Learning Center (OLC) at www.mhhe.com/zoology Vertebrates: Macroscopic Anatomy of the Digestive System
(click on the book’s title) to find the following chapter-related materials: Vertebrates: Microscopic Anatomy of the Digestive System
• CHAPTER QUIZZING
Nutrition
• ANIMATIONS
Human Digestive Topics
Digestion • BOXED READINGS ON
Plaque Formation Suspension Feeding in Invertebrates and Nonvertebrate Chordates
Dental Caries Filter Feeding in Vertebrates
Endoscopy • SUGGESTED READINGS
Digestion (Mouth to Stomach) • LAB CORRELATIONS
Digestion (Stomach) Check out the OLC to find specific information on these related lab
Stomach Digestion exercises in the General Zoology Laboratory Manual, 5th edition, by
Ulcers Stephen A. Miller:
Formation of Gallstones
Small Intestine Digestion Exercise 22 Vertebrate Digestion
Digestion (Stomach to Small Intestine)
Miller−Harley: Zoology, III. Form and Function: A 28. Temperature and Body © The McGraw−Hill
Fifth Edition Comparative Perspecitive Fluid Regulation Companies, 2001
CHAPTER 28
T E M P E R AT U R E A N D B O D Y F L U I D R E G U L AT I O N
Outline Concepts
Homeostasis and Temperature Regulation 1. Thermoregulation is a complex and important physiological process that maintains, to
The Impact of Temperature on varying degrees, an animal’s body temperature, despite variations in environmental tem-
Animal Life perature. Based on this regulation, animals can be categorized as endotherms or
Heat Gains and Losses ectotherms, and homeotherms or heterotherms.
Some Solutions to Temperature 2. For osmoregulation, some invertebrates have contractile vacuoles, flame-cell systems,
Fluctuations antennal (green) glands, maxillary glands, coxal glands, nephridia, or Malpighian
Temperature Regulation in Invertebrates tubules.
Temperature Regulation in Fishes 3. A vertebrate’s urinary system functions in osmoregulation and excretion, both of which
Temperature Regulation in Amphibians
are necessary for internal homeostasis. Osmoregulation governs water and salt balance,
and Reptiles
and excretion eliminates metabolic wastes. In fishes, reptiles, birds, and mammals, the
Temperature Regulation in Birds and
kidneys are the primary osmoregulatory structures.
Mammals
Heat Production in Birds and Mammals
The earth’s environments vary dramatically in temperature and amount of water present.
Control of Water and Solutes
In the polar regions, high mountain ranges, and deep oceans, the temperature remains
(Osmoregulation and Excretion)
near or below 0° C (32° F) throughout the year. Temperatures exceeding 40° C (103° F)
Invertebrate Excretory Systems
are common in equatorial deserts. Between these two extremes, in the earth’s temperate
Contractile Vacuoles
regions, temperatures fluctuate widely. The temperate regions have varying amounts of
Protonephridia
water, as well as varied habitats—freshwater, saltwater, wetlands, mountains, and
Metanephridia
grasslands.
Antennal (Green) and Maxillary
Animals have successfully colonized these varied places on earth by possessing
Glands
homeostatic mechanisms for maintaining a relatively constant internal environment,
Malpighian Tubules
despite fluctuations in the external environment. This chapter covers three separate but
Coxal Glands
related homeostatic systems that enable animals to survive the variations in temperature,
Vertebrate Excretory Systems
water availability, and salinity (salt concentration) on the earth. The thermoregulatory
How Vertebrates Achieve
system maintains an animal’s body temperature and/or its responses to shifts in environ-
Osmoregulation
mental temperature. The osmoregulatory system maintains the level and concentration of
Vertebrate Kidney Variations
water and salts in the body. And the urinary system eliminates metabolic wastes from the
How the Metanephric Kidney
body and functions in osmoregulation.
Functions
HOMEOSTASIS AND TEMPERATURE

REGULATION
The temperature of a living cell affects the rate of its metabolic processes. An animal can
grow faster and respond to the environment more rapidly if its cells are kept warm.
In fact, zoologists believe that the ability of some higher animals to maintain a
This chapter contains evolutionary concepts, which are set off in this font.
455
constant (homeostatic), relatively high body temperature is a

major reason for their evolutionary success. This ability to con-
trol the temperature of the body is called thermoregulation
(“heat control”) and involves the nervous, endocrine, respiratory,
and circulatory systems in higher animals. Evaporation
from the mouth
THE IMPACT OF TEMPERATURE Convection

ON ANIMAL LIFE Radiation
Every animal’s physiological functions are inexorably linked to

temperature, because metabolism is sensitive to changes in inter-
nal temperature. Thus, temperature has been a strong source of
selective pressure on all animals. The rate of cellular respiration
increases with temperature up to a point. When the temperature
rises above the temperature optima at which enzymes most effi-
ciently catalyze their chemical reactions, the rates decline as the Hot rock
Conduction
enzymes begin to denature. The chemical interactions holding the
enzymes in their three-dimensional shape are also disrupted. FIGURE 28.1
Thus, the results of enzyme evolution have frequently been Heat Gain and Loss for a Terrestrial Reptile in a Typical Terrestrial
enzymes with temperature optima that reflect an animal’s habi- Environment. Heat is either gained or lost by objects in direct contact
tat. For example, a digestive enzyme in a trout might function with the animal (conduction), by air currents (convection), by exhaled
optimally at 10° C, whereas another enzyme in the human body air (evaporation), or by electromagnetic waves (radiation).
that catalyzes the same reaction functions best at 37° C. Higher
temperatures cause the proteins in nucleic acids to denature, and
lower temperatures may cause membranes to change from a fluid Evaporation is loss of heat from a surface as water molecules
to a solid state, which can interfere with many cellular processes, escape in the form of a gas. It is useful only to terrestrial animals.
such as active-transport pumps. For example, humans and some other mammals (chimpanzees and
Animals can guard against these damaging effects of tem- horses) have sweat glands that actively move watery solutions
perature fluctuations by balancing heat gains and heat losses with through pores to the skin surface. When skin temperature is high,
their environment. water at the surface absorbs enough thermal energy to break the
hydrogen bonds holding the individual water molecules together,
HEAT GAINS AND LOSSES and they depart from the surface, carrying heat with them. As
long as the environmental humidity is low enough to permit com-
Animals produce heat as a by-product of metabolism and either plete evaporation, sweating can rid the mammalian body of excess
gain heat from, or lose it to, the environment. The total body tem- heat; however, the water must evaporate. Sweat dripping from a
perature is a result of an interaction of these factors and can be mammal has no cooling effect at all.
expressed as: Radiation is the emission of electromagnetic waves that
objects, such as another animal’s body or the sun, produce. Radi-
Body temperature heat produced metabolically ation can transfer heat between objects that are not in direct
heat gained from the environment contact with each other, as happens when an animal suns itself
heat lost to the environment (figure 28.2).
Animals use four physical processes to exchange heat with the
environment: conduction, convection, evaporation, and radiation
(figure 28.1). Conduction is the direct transfer of thermal motion SOME SOLUTIONS TO
(heat) between molecules of the environment and those on the body TEMPERATURE FLUCTUATIONS
surface of an animal. This transfer is always from an area of higher
temperature to one of lower temperature because heat moves down Animals cope with temperature fluctuations in one of three basic
thermal gradients. For example, when you sit on the cold ground, ways. (1) They can occupy a place in the environment where the
you lose heat, and when you sit on warm sand, you gain heat. temperature remains constant and compatible with their physio-
Convection is the movement of air (or a liquid) over the logical processes; (2) their physiological processes may have
surface of a body; it contributes to heat loss if the air is cooler than adapted to the range of temperatures in which the animals are
the body or heat gain if the air is warmer than the body. On a cool capable of living; or (3) they can generate and trap heat internally
day, your body loses heat by convection because your skin temper- to maintain a constant body temperature, despite fluctuations in
ature is higher than the surrounding air temperature. the temperature of the external environment.
CHAPTER 28 Temperature and Body Fluid Regulation 457
(have a variable body temperature), there are many exceptions.

Some endotherms vary their body temperatures seasonally (e.g.,
hibernation); others vary it on a daily basis.
For example, some birds (e.g., hummingbirds) and mam-
mals (e.g., shrews) can only maintain a high body temperature for
a short period because they usually weigh less than 10 g and have
a body mass so small that they cannot generate enough heat to
compensate for the heat lost across their relatively large surface
area. Hummingbirds must devote much of the day to locating and
sipping nectar (a very high-calorie food source) as a constant
energy source for metabolism. When not feeding, hummingbirds
rapidly run out of energy unless their metabolic rates decrease
considerably. At night, hummingbirds enter a sleep-like state,
called daily torpor, and their body temperature approaches that of
the cooler surroundings. Some bats also undergo daily torpor to
conserve energy.
Some ectotherms can maintain fairly constant body temper-
atures. Among these are a number of reptiles that can maintain
fairly constant body temperatures by changing position and loca-
tion during the day to equalize heat gain and loss.
In general, ectotherms are more common in the tropics
because they do not have to expend as much energy to maintain
body temperature there, and they can devote more energy to food
FIGURE 28.2 gathering and reproduction. Indeed, in the tropics, amphibians
Radiation Warms an Animal. After a cold night in its den on the are far more abundant than mammals. Conversely, in moderate
Kalahari Desert, a meerkat (Suricata suricatta) stands at attention, allow-
ing the large surface area of its body to absorb radiation from the sun.
to cool environments, endotherms have a selective advantage
and are more abundant. Their high metabolic rates and insula-
tion allow them to occupy even the polar regions (e.g., polar
Animals can be categorized as ectotherms or endotherms, bears). In fact, the efficient circulatory systems of birds and mam-
based on whether their source of body heat is from internal mals can be thought of as adaptations to endothermy and a high
processes or derived from the environment. Ectotherms (Gr. metabolic rate.
ectos, outside) derive most of their body heat from the environ-
ment rather than from their own metabolism (figure 28.3). They
have low rates of metabolism and are poorly insulated. In general, TEMPERATURE REGULATION
reptiles, amphibians, fishes, and invertebrates are ectotherms, IN INVERTEBRATES
although a few reptiles, insects, and fishes can raise their internal
temperature. Ectotherms tend to move about the environment As previously noted, environmental temperature is critical in lim-
and find places that minimize heat or cold stress to their bodies. iting the distribution of all animals and in controlling metabolic
Birds and mammals are called endotherms (Gr. endos, reactions. Many invertebrates have relatively low metabolic rates
within) because they obtain their body heat from cellular and have no thermoregulatory mechanisms; thus, they passively
processes. A constant source of internal heat allows them to main- conform to the temperature of their external environment. These
tain a nearly constant core temperature, despite the fluctuating invertebrates are termed thermoconformers.
environmental temperature. (“Core” refers to the body’s internal Evidence indicates that some higher invertebrates can
temperature as opposed to the temperature near its surface.) directly sense differences in environmental temperatures; how-
Most endotherms have bodies insulated by fur or feathers ever, specific receptors are either absent or unidentified. What
and a relatively large amount of fat. This insulation enables them zoologists do know is that many arthropods, such as insects, crus-
to retain heat more efficiently and to maintain a high core tem- taceans, and the horseshoe crab (Limulus), can sense thermal vari-
perature. Endothermy allows animals to stabilize their core tem- ation. For example, ticks of warm-blooded vertebrates can sense
perature so that biochemical processes and nervous system func- the “warmth of a nearby meal” and drop on the vertebrate host.
tions can proceed at steady, high levels. Endothermy allows some Many arthropods have unique mechanisms for surviving
animals to colonize habitats denied to ectotherms. temperature extremes. For example, temperate-zone insects avoid
Another way of categorizing animals is based on whether freezing by reducing the water content in their tissues as winter
they maintain a constant or variable body temperature. Although approaches. Other insects can produce glycerol or other glycopro-
most endotherms are homeotherms (maintain a relatively con- teins that act as an antifreeze. Some moths and bumblebees warm
stant body temperature), and most ectotherms are heterotherms up prior to flight by shivering contractions of their thoracic flight
FIGURE 28.3
Heat Gain in an Insect. Postures a dragonfly adopts to either maximize or minimize heat gain.
muscles. Most large, flying insects have evolved a mechanism to duced sufficient metabolic heat that thermoregulatory strate-
prevent overheating during flight; blood circulating through the gies could evolve. An increased locomotor metabolism could
flight muscles carries heat from the thorax to the abdomen, well have preceded the evolution of thermoregulation in
which gets rid of the heat—much as coolant circulating through vertebrates.
an automobile engine passes through the radiator. Certain
cicadas (Diceroprocta apache) that live in the Sonoran Desert
have independently evolved the complete repertoire of evapo- TEMPERATURE REGULATION IN FISHES
rative cooling mechanisms that vertebrates use. When threat-
ened with overheating, these cicadas extract water from their The temperature of the surrounding water determines the body
blood and transport it through large ducts to the surface of their temperature of most fishes. Fishes that live in extremely cold
body, where it passes through sweat pores and evaporates. In water have “antifreeze” materials in their blood. Polyalcohols
other words, these insects can sweat. (e.g., sorbitol, glycerol) or water-soluble peptides and glycopep-
Body posture and orientation of the wings to the sun can tides lower the freezing point of blood plasma and other body flu-
markedly affect the body temperature of basking insects. For ids. These fishes also have proteins or protein-sugar compounds
example, perching dragonflies and butterflies can regulate their that stunt the growth of ice crystals that begin to form. These
radiation heat gain by postural adjustments (figure 28.3). adaptations enable these fishes to stay flexible and swim freely in a
To prevent overheating, many ground-dwelling arthropods supercooled state (i.e., at a temperature below the normal freezing
(Tenebrio beetles, locusts, scorpions) raise their bodies as high off temperature of a solution).
the ground as possible to minimize heat gain from the ground. Some active fishes maintain a core temperature significantly
Some caterpillars and locusts orient with reference to both the sun above the temperature of the water. Bluefin tuna and the great
and wind to vary both radiation heat gain and convective heat white shark have major blood vessels just under the skin. Branches
loss. Some desert-dwelling beetles can exude waxes from thou- deliver blood to the deeper, powerful, red swimming muscles, where
sands of tiny pores on their cuticle. These “wax blooms” prevent smaller vessels are arranged in a countercurrent heat exchanger
dehydration and also are an extra barrier against the desert sun. called the rete mirabile (“miraculous net”) (figure 28.4). The heat
Color has a significant effect on thermoregulation since that these red muscles generate is not lost because it is transferred in
50% of the radiant energy from the sun is in the visible spectrum. the rete mirabile from venous blood passing outward to cold arterial
A black surface reflects less radiant energy than a white surface. blood passing inward from the body surface. This arrangement of
Thus, many black beetles may be more active earlier in the day blood vessels enhances vigorous activity by keeping the swimming
because they absorb more radiation and heat faster. Conversely, muscles several degrees warmer than the tissue near the surface of
white beetles are more active in the hotter parts of the day because the fish. This system has been adaptive for these fishes. Their mus-
they absorb less heat. cular contractions can have four times as much power as those of
The previous examples of invertebrate temperature reg- similar muscles in fishes with cooler bodies. Thus, they can swim
ulation give clues to how thermoregulation may have evolved faster and range more widely through various depths in search of
in vertebrates. The endothermic temperature regulation of prey than can other predatory fishes more limited to given water
active insects apparently evolved because locomotion pro- depths and temperatures.
FIGURE 28.4
Thermoregulation in Large, Active Fishes. In the bluefin tuna, the rete mirabile of arteries and veins acts as a countercurrent exchange system that
helps reduce the loss of body heat. The cross section through the body shows that the temperature is highest around the red swimming muscles.
TEMPERATURE REGULATION also have an expandable rib cage, which allows for more powerful
IN AMPHIBIANS AND REPTILES and efficient ventilation. Reptiles are almost completely ectother-
mic. They have a low metabolic rate and warm themselves by
Animals, such as amphibians and reptiles, that have air rather behavioral adaptations. In addition, some of the more sophisti-
than water as a surrounding medium face marked daily and sea- cated regulatory mechanisms found in mammals are first found in
sonal temperature changes. Most of these animals are ectotherms. reptiles. For example, diving reptiles (e.g., sea turtles, sea snakes)
They derive heat from their environment, and their body temper- conserve body heat by routing blood through circulatory shunts
atures vary with external temperatures. into the center of the body. These animals can also increase heat
Most amphibians have difficulty in controlling body heat production in response to the hormones thyroxine and epineph-
because they produce little of it metabolically and rapidly lose rine. In addition, tortoises and land turtles can cool themselves
most of it from their body surfaces. However, as previously noted, through salivating and frothing at the mouth, urinating on the
behavioral adaptations enable them to maintain their body tem- back legs, moistening the eyes, and panting.
perature within a homeostatic range most of the time. Amphib-
ians have an additional thermoregulatory problem because they
must exchange oxygen and carbon dioxide across their skin sur- TEMPERATURE REGULATION
face, and this moisture layer acts as a natural evaporative cooling IN BIRDS AND MAMMALS
system. This problem of heat loss through evaporation limits the
habitats and activities of amphibians to warm, moist areas. Some Birds and mammals are the most active and behaviorally complex
amphibians, such as bullfrogs, can vary the amount of mucus they vertebrates. They can live in habitats all over the earth because
secrete from their body surface—a physiological response that they are homeothermic endotherms; they can maintain body tem-
helps regulate evaporative cooling. peratures between 35 and 42° C with metabolic heat.
Reptiles have dry rather than moist skin, which reduces the Various cooling mechanisms prevent excessive warming in
loss of body heat through evaporative cooling of the skin. They birds. Because they have no sweat glands, birds pant to lose heat
through evaporative cooling. Some species have a highly vascu-

larized pouch (gular pouch) in their throat that they can flutter (a
process called gular flutter) to increase evaporation from the res-
piratory system.
Some birds possess mechanisms for preventing heat loss.
Feathers are excellent insulators for the body, especially downy-
type feathers that trap a layer of air next to the body to reduce
heat loss from the skin (figure 28.5a). (This mechanism explains
why goose down is such an excellent insulator and is used in out-
door vests and coats for protection from extreme cold.) Aquatic
species, who lose heat from their legs and feet, have peripheral
countercurrent heat exchange vessels called a rete mirabile in
their legs to reduce heat loss (figure 28.5b). Mammals that live in
cold regions, such as the arctic fox and barren-ground caribou,
also have these exchange vessels in their extremities (e.g., legs,
tails, ears, nose). Animals in hot climates, such as jackrabbits, (a)
have mechanisms (e.g., large ears) to rid the body of excess heat
Venous Arterial
(figure 28.6). blood blood
Thick pelts and a thick layer of insulating fat called blubber 33° 35°C
just under the skin help marine animals, such as seals and whales,
to maintain a body temperature of around 36 to 38° C. In the tail
27° 30°
and flippers, which have no blubber, a countercurrent system of
arteries and veins helps minimize heat loss.
Birds and mammals also use behavioral mechanisms to cope 18° 20°
with external temperature changes. Like ectotherms, they sun
themselves or seek shade as the temperature fluctuates. Many ani-
mals huddle to keep warm; others share burrows for protection 9° 10°
from temperature extremes. Migration to warm climates and
hibernation enable many different birds and mammals to survive
the harsh winter months. Others, such as the desert camel, have a
multitude of evolutionary adaptations for surviving in some of the
hottest and driest climates on earth.
HEAT PRODUCTION IN BIRDS (b)
AND MAMMALS FIGURE 28.5

Insulation and Countercurrent Heat Exchange. (a) A thick layer of
In endotherms, heat generation can warm the body as it dissipates down feathers keeps these Chinstrap penguins (Pygocelis antarctia)
throughout tissues and organs. Birds and mammals can generate warm. Their covering of short, stiff feathers interlocks to trap air, form-
heat (thermogenesis) by muscle contraction, ATPase pump ing the ornithological equivalent of a diver’s suit. (b) The countercur-
enzymes, oxidation of fatty acids in brown fat, and other meta- rent heat exchanger in a bird foot. Some aquatic birds, such as this duck,
possess countercurrent systems of arteries and veins (rete mirabile) in
bolic processes. their legs that reduce heat loss. The arteries carry warm blood down the
Every time a muscle cell contracts, the actin and myosin fil- legs to warm the cooler blood in the veins, so that the heat is carried
aments sliding over each other and the hydrolysis of ATP mole- back to the body rather than lost through the feet that are in contact
cules generate heat. Both voluntary muscular work (e.g., running, with a cold surface.
flying, jumping) and involuntary muscular work (e.g., shivering)
generate heat. Heat generation by shivering is called shivering
thermogenesis. releasing heat energy. The hormonal triggering of heat production
Birds and mammals have a unique capacity to generate heat is called nonshivering thermogenesis.
by using specific enzymes of ancient evolutionary origin—the Brown fat is a specialized type of fat found in newborn
ATPase pump enzymes in the plasma membranes of most cells. mammals, in mammals that live in cold climates, and in mammals
When the body cools, the thyroid gland releases the hormone thy- that hibernate (figure 28.7). The brown color of this fat comes
roxine. Thyroxine increases the permeability of many cells to from the large number of mitochondria with their iron-containing
sodium (Na) ions, which leak into the cells. The ATPase pump cytochromes. Deposits of brown fat are beneath the ribs and in the
quickly pumps these ions out. In the process, ATP is hydrolyzed, shoulders. A large amount of heat is produced when brown fat
Brown fat
(a) Brown fat
Warm blood
FIGURE 28.6
Temperature Regulation. This antelope jackrabbit (Lepus alleni) must
get rid of excess body heat. Its huge, thin, highly vascularized ears have a
large surface area for heat exchange.
cells oxidize fatty acids, because little ATP is made. Blood flowing
past brown fat is heated and contributes to warming the body. (b) Warmer blood
The basal metabolic rate of birds and mammals is high and
also produces heat as an inadvertent but useful by-product. FIGURE 28.7
In amphibians, reptiles, birds, and mammals, specialized Brown Fat. (a) Many mammals, such as this bat, have adipose tissue
cells in the hypothalamus of the brain control thermoregulation. called brown fat between the shoulder blades. (b) The area of brown fat
is much warmer than the rest of the body. Blood flowing through the
The two hypothalamic thermoregulatory areas are the heating brown fat is warmed.
center and the cooling center. The heating center controls vaso-
constriction of superficial blood vessels, erection of hair and fur,
and shivering or nonshivering thermogenesis. The cooling center
controls vasodilation of blood vessels, sweating, and panting. CONTROL OF WATER AND
Overall, feedback mechanisms (with the hypothalamus acting as a SOLUTES (OSMOREGULATION
thermostat) trigger either the heating or cooling of the body and AND EXCRETION)
thereby control body temperature (figure 28.8). Specialized neu-
ronal receptors in the skin and other parts of the body sense tem- Excretion (L. excretio, to eliminate) can be defined broadly as the
perature changes. Warm neuronal receptors excite the cooling elimination of metabolic waste products from an animal’s body.
center and inhibit the heating center. Cold neuronal receptors These products include carbon dioxide and water (which cellular
have the opposite effects. respiration primarily produces), excess nitrogen (which is pro-
During the winter, various endotherms (e.g., bats, wood- duced as ammonia from deamination of amino acids), and solutes
chucks, chipmunks, ground squirrels) go into hibernation (L. (various ions). Chapter 26 covers the excretion of respiratory car-
hiberna, winter). During hibernation, the metabolic rate slows, as bon dioxide.
do the heart and breathing rates. Mammals prepare for hiberna- Excretion of nitrogenous wastes is usually associated with
tion by building up fat reserves and growing long winter pelts. All the regulation of water and solute (ionic) balance by a physiolog-
hibernating animals have brown fat. Decreasing day length stimu- ical process called osmoregulation. Osmoregulation is necessary
lates both increased fat deposition and fur growth. Another phys- for animals in all habitats. If the osmotic concentration of the
iological state characterized by slow metabolism and inactivity is body fluids of an animal equals that of the medium (the animal’s
aestivation (L. aestivus, summer), which allows certain mammals environment), the animal is an osmoconformer. When the
to survive long periods of elevated temperature and diminished osmotic concentration of the environment changes, so does that
water supplies. of the animal’s body fluids. Obviously, this type of osmoregulation
Some animals, such as badgers, bears, opossums, raccoons, is not efficient and has limited the distribution of those animals
and skunks, enter a state of prolonged sleep in the winter. Since using it. In contrast, an animal that maintains its body fluids at a
their body temperature remains near normal, this is not true different osmotic concentration from that of its surrounding envi-
hibernation. ronment is an osmoregulator.
Changes in Changes in
environmental body (blood)
temperature temperature
Thermoreceptors Thermoreceptors
found in the found inside
skin the body
BRAIN HYPOTHALAMUS
Thyroid-releasing hormone
Autonomic
Motor area Anterior
nervous system
of cerebral pituitary gland
(sympathetic or
cortex (adenohypophysis)
parasympathetic)
Thyroid-stimulating
hormone (thyrotropin)
Voluntary Involuntary Thyroid

response responses gland
Thyroxine
Skeletal Blood Adrenal glands Body cells Sweat glands

muscles vessels (cortex)
Epinephrine
Shivering Vasoconstriction Increases or Basal metabolic Sweating cools

thermogenesis (sympathetic) or decreases rate increases the body (inhibition
(generates vasodilation organic or decreases retains heat)
heat) (parasympathetic) metabolism
Either heat generation or conservation
Blood temperature either rises or falls
FIGURE 28.8
Thermoregulation. Overview of the feedback pathways that control the core body temperature of a mammal. Arrows show the major control pathways.
Animals living in seawater have body fluids with an osmotic water and prevent fluid accumulation. Land animals have a higher
concentration that is about a third less (hypoosmotic) than the sur- concentration of water in their fluids than in the surrounding air.
rounding seawater, and water tends to leave their bodies continu- They tend to lose water to the air through evaporation and may use
ally. To compensate for this problem, mechanisms evolved in considerable amounts of water to dispose of wastes.
these animals to conserve water and prevent dehydration. Fresh- The form and function of organs or systems associated with
water animals have body fluids that are hyperosmotic with excretion and osmoregulation are related both to environmental
respect to their environment, and water tends to continually enter conditions (saltwater, freshwater, terrestrial) and to body size
their bodies. Mechanisms evolved in these animals that excrete (especially the surface-to-volume ratio).
Flame cell
Cilia forming
Nucleus “flame”
Fenestration
Path of fluid
Nephridiopore
Flame cell Cell of

tubule wall
Excretory Tubule
tube lumen
To nephridiopore
(b)
FIGURE 28.9
Contractile Vacuoles. A photomicrograph (100) showing the location
of two contractile vacuoles (black arrows) in a stained Paramecium. Notice (a)
the small tubules surrounding each vacuole. These tubes collect water and
deliver it to the contractile vacuole, which expels the fluid through a pore. FIGURE 28.10
Protonephridial (Excretory) System in a Turbellarian. (a) The sys-
tem lies in the mesenchyme and consists of a network of fine tubules
INVERTEBRATE EXCRETORY that run the length of the animal on each side and open to the surface
SYSTEMS by minute excretory pores called nephridiopores. (b) Numerous fine side
branches from the tubules originate in the mesenchyme in enlargements
Aquatic invertebrates occur in a wide range of media, from fresh- called flame cells.
water to markedly hypersaline water (e.g., salt lakes). Generally,
marine invertebrates have about the same osmotic concentration
as seawater (i.e., they are osmoconformers). This avoids any need PROTONEPHRIDIA
to osmoregulate. Most water and ions are gained across the integu-
ment, via gills, by drinking, and in food. Ions and wastes are Although a few groups of metazoan invertebrates possess no
mostly lost by diffusion via the integument, gills, or urine. known excretory structures, most have nephridia (Gr. nephros,
Freshwater invertebrates are strong osmoregulators because kidney) (sing., nephridium) that serve for excretion, osmoregula-
it is impossible to be isosmotic with dilute media. Any water gain tion, or both. Probably the earliest type of nephridium to
is usually eliminated as urine. appear in the evolution of animals was the protonephrid-
A number of invertebrate taxa have more or less successfully ium (Gr. protos, first nephridium).
invaded terrestrial habitats. The most successful terrestrial inver- Among the simplest of the protonephridia are flame-cell
tebrates are the arthropods, particularly the insects, spiders, scor- systems, such as those in rotifers, some annelids, larval molluscs,
pions, ticks, mites, centipedes, and millipedes. Overall, the water and some flatworms (figure 28.10). The protonephridial excretory
and ion balance of terrestrial invertebrates is quite different from system is composed of a network of excretory canals that open to
that of aquatic animals because terrestrial invertebrates face lim- the outside of the body through excretory pores. Bulblike flame
ited water supplies and water loss by evaporation from their cells are located along the excretory canals. Fluid filters into the
integument. Some of the invertebrate excretory mechanisms and flame cells from the surrounding interstitial fluid, and beating cilia
systems are now discussed. propel the fluid through the excretory canals and out of the body
through the excretory pores. Flame-cell systems function prima-
rily in eliminating excess water. Nitrogenous waste simply diffuses
CONTRACTILE VACUOLES across the body surface into the surrounding water.
Some protists and marine invertebrates (e.g., protozoa, cnidarians,
echinoderms, sponges) do not have specialized excretory structures METANEPHRIDIA
because wastes simply diffuse into the surrounding isoosmotic
water. In some freshwater species, cells on the body surface A more advanced type of excretory structure among invertebrates
actively pump ions into the animal. Many freshwater species (pro- is the metanephridium (Gr. meta, beyond nephridium;) (pl.,
tozoa, sponges), however, have contractile vacuoles that pump out metanephridia). Protonephridia and metanephridia have critical
excess water. Contractile vacuoles are energy-requiring devices structural differences. Both open to the outside, but meta-
that expel excess water from individual cells exposed to hypoos- nephridia (1) also open internally to the body fluids and (2) are
motic environments (figure 28.9). multicellular.
Intestine Capillary network Septum Bladder

Antennal (green) gland
Nephridial
canal
Excretory
pore
Nephrostome Tubule Bladder Nephridiopore
Anterior Posterior
FIGURE 28.11
Earthworm Metanephridium. The metanephridium opens by a cili-
ated nephrostome into the cavity of one segment, and the next segment
contains the nephridiopore. The main tubular portion of the
metanephridium is coiled and is surrounded by a capillary network. Bladder
Waste can be stored in a bladder before being expelled to the outside.
Most segments contain two metanephridia. End sac Labyrinth
FIGURE 28.12
Antennal (Green) Gland of the Crayfish. The antennal gland, which
Most annelids (such as the common earthworm) and a vari- lies in front of and to both sides of the esophagus, is divided into an end
ety of other invertebrates have a metanephridial excretory system. sac, where fluid collects by filtration, and a labyrinth. The labyrinth
Recall that the earthworm’s body is divided into segments and walls are greatly folded and glandular and appear to be an important site
for reabsorption. The labyrinth leads via a nephridial canal into a blad-
that each segment has a pair of metanephridia. Each meta-
der. From the bladder, a short duct leads to an excretory pore.
nephridium begins with a ciliated funnel, the nephrostome, that
opens from the body cavity of a segment into a coiled tubule (fig-
ure 28.11). As beating cilia move the fluid through the tubule, a
network of capillaries surrounding the tubule reabsorbs and car- In other crustaceans (some malacostracans [crabs, shrimp,
ries away ions. Each tubule leads to an enlarged bladder that emp- pillbugs]), the excretory organs are near the maxillary segments
ties to the outside of the body through an opening called the and are termed maxillary glands. In maxillary glands, fluid col-
nephridiopore. lects within the tubules from the surrounding blood of the hemo-
The excretory system of molluscs includes protonephridia in coel, and this primary urine is modified substantially by selective
larval stages and metanephridia in adults. reabsorption and secretion as it moves through the excretory sys-
tem and rectum.
ANTENNAL (GREEN) AND

MAXILLARY GLANDS MALPIGHIAN TUBULES
Insects have an excretory system made up of the gut and
In those crustaceans that have gills, nitrogenous wastes are
Malpighian tubules attached to the gut (figure 28.13). Excretion
removed by simple diffusion across the gills. Most crustaceans
involves the active transport of potassium ions into the tubules
release ammonia, although they also produce some urea and uric
from the surrounding hemolymph and the osmotic movement of
acid as waste products. Thus, the excretory organs of freshwater
water that follows. Nitrogenous waste (uric acid) also enters the
species may be more involved with the reabsorption of ions and
tubules. As fluid moves through the Malpighian tubules, some of
elimination of water than with the discharge of nitrogenous
the water and certain ions are recovered. All of the uric acid
wastes. The excretory organs in some crustaceans (crayfish,
passes into the gut and out of the body.
crabs) are antennal glands or green glands because of their loca-
tion near the antenna and their green color (figure 28.12). Fluid
filters into the antennal gland from the hemocoel. Hemolymph COXAL GLANDS
pressure from the heart is the main driving force for filtration.
Marine crustaceans have a short nephridial canal and produce Coxal (L. coxa, hip) glands are common among arachnids (spi-
urine that is isoosmotic to their hemolymph. The nephridial canal ders, scorpions, ticks, mites). These spherical sacs resemble
is longer in freshwater crustaceans, which allows more surface area annelid nephridia (figure 28.14). Wastes are collected from the
for ion transport. surrounding hemolymph of the hemocoel and discharged through
Active transport
Stercoral
Passive movement
pocket
Defecation
Water/ Na+K+ Uric

Midgut sugars/ acid
amino acids
Malpighian
tubule
Anus
Coxal Midgut Malpighian
glands tubule
(a)
Hindgut Ventral
Dorsal dilation
dilation
Water
Na+/K+/sugars/amino acids
Accessory
Rectum gland
Uric acid
FIGURE 28.13
Malpighian Tubules. Malpighian tubules remove nitrogenous wastes
(uric acid) from the hemocoel. Various ions are actively transported across Duct
(b) Filtration
the outer membrane of the tubule. Water follows these ions into the membrane Coxal gland
tubule and carries amino acids, sugars, and some nitrogenous wastes along muscles
passively. Some water, ions, and organic compounds are reabsorbed in the
basal portion of the Malpighian tubules and the hindgut; the rest are reab- FIGURE 28.14
sorbed in the rectum. Uric acid moves into the hindgut and is excreted.
Coxal Glands in Arachnids. (a) The gut and excretory systems of a
spider. (b) Coxal gland muscles attach to the thin saccular filtration
membrane. These muscles promote filtration and fluid flow (black ar-
pores on from one to several pairs of appendages near the proximal rows) by contracting and relaxing along the tubular duct. Water and
joint (coxa) of the leg. Recent evidence suggests that the coxal solutes are reabsorbed along the tubular duct.
glands may also function in the release of pheromones.
Other arachnid species have Malpighian tubules instead of,
or in addition to, the coxal glands. In some of these species, how- Vertebrates live in saltwater, freshwater, and on land; each
ever, Malpighian tubules seem to function in silk production of these environments presents different water and solute prob-
rather than in excretion. lems that vertebrates have solved in different ways. The next sec-
tion discusses how vertebrates avoid losing or gaining too much
water and, in turn, how they maintain a homeostatic solute con-
VERTEBRATE EXCRETORY centration in their body fluids. The disposal (excretion) of certain
metabolic waste products is also coupled with osmotic balance
SYSTEMS
and is discussed with the urinary system.
Vertebrates face the same problems as invertebrates in controlling
water and ion balance. Generally, water losses are balanced pre-
cisely by water gains (table 28.1). Vertebrates gain water by HOW VERTEBRATES ACHIEVE
absorption from liquids and solid foods in the small and large OSMOREGULATION
intestines, and by metabolic reactions that yield water as an end
A variety of mechanisms have evolved in vertebrates to cope with
product. They lose water by evaporation from respiratory surfaces,
their osmoregulatory problems, and most of them are adaptations
evaporation from the integument, sweating or panting, elimina-
of the urinary system. As presented in chapter 26, vertebrates
tion in feces, and excretion by the urinary system.
have a closed circulatory system containing blood that is under
Solute losses also must be balanced by solute gains. Verte-
pressure. This pressure forces blood through a membrane filter in a
brates take in solutes by absorption of minerals from the small and
kidney, where the following three key functions take place:
large intestines, through the integument or gills, from secretions
of various glands or gills, and by metabolism (e.g., the waste prod- 1. Filtration, in which blood passes through a filter that retains
ucts of degradative reactions). They lose solutes in sweat, feces, blood cells, proteins, and other large solutes but lets small
urine, and gill secretions, and as metabolic wastes. The major molecules, ions, and urea pass through
metabolic wastes that must be eliminated are ammonia, urea, or 2. Reabsorption, in which selective ions and molecules are
uric acid. taken back into the bloodstream from the filtrate
TA B L E 2 8 . 1
AVERAGE WATER GAIN AND LOSS IN A HUMAN AND KANGAROO RAT
VERTEBRATE WATER GAIN (ML) WATER LOSS (ML)
Human (daily) Ingested in solid food 1,200 Feces 100

Ingested as liquids 1,000 Urine 1,500
Metabolically produced 350 Skin and lungs 950
Total 2,550 2,550
Kangaroo rat (over 4 weeks) Ingested in solid food 6 Feces 3

Ingested in liquids 0 Urine 13
Metabolically produced 54 Skin and lungs 44
Total 60 60
3. Secretion, whereby select ions and end products of metabo- Sharks

lism (e.g., K, H, NH3) that are in the blood are added to
Sharks and their relatives (skates and rays) have mesonephric kid-
the filtrate for removal from the body
neys and have solved their osmotic problem in ways different from
the bony fishes (figure 28.15b). Instead of actively pumping ions
out of their bodies through the kidneys, they have a rectal gland
VERTEBRATE KIDNEY VARIATIONS that secretes a highly concentrated salt (NaCl) solution. To
reduce water loss, they use two organic molecules—urea and
Vertebrates have two kidneys that are in the back of the abdomi-
trimethylamine oxide (TMO)—in their body fluids to raise the
nal cavity, on either side of the aorta. Each kidney has a coat of
osmotic pressure to a level equal to or higher than that of the
connective tissue called the renal capsule (L. renes, kidney). The
seawater.
inner portion of the kidney is called the medulla; the region
Urea denatures proteins and inhibits enzymes, whereas
between the capsule and the medulla is the cortex.
TMO stabilizes proteins and activates enzymes. Together in the
The structure and function of vertebrate kidneys differ,
proper ratio, they counteract each other, raise the osmotic pres-
depending on the vertebrate groups and the developmental stage.
sure, and do not interfere with enzymes or proteins. This reciproc-
Overall, there are three kinds of vertebrate kidneys: the
ity is termed the counteracting osmolyte strategy.
pronephros, mesonephros, and metanephros. The pronephros
A number of other fishes and invertebrates have evolved
appears only briefly in many vertebrate embryos, and not at all in
the same mechanism and employ pairs of counteracting
mammalian embryos (figure 28.15a). In some vertebrates, the
osmolytes to raise the osmotic pressure of their body fluids.
pronephros is the first osmoregulatory and excretory organ of the
embryo (tadpoles and other amphibian larvae); in others (hag-
fishes), it remains as the functioning kidney. During embryonic Teleost Fishes
development of amniotes, or during metamorphosis in amphib- Most teleost fishes have mesonephric kidneys. Because the body
ians, the mesonephros replaces the pronephros (figure 28.15b). fluids of freshwater fishes are hyperosmotic relative to freshwater
The mesonephros is the functioning embryonic kidney of many (see table 28.2), water tends to enter the fishes, causing excessive
vertebrates and also adult fishes and amphibians. The mesonephros hydration or bloating (figure 28.16a). At the same time, body ions
gives way during embryonic development to the metanephros in tend to move outward into the water. To solve this problem, fresh-
adult reptiles, birds, and mammals (figure 28.15c). water fishes usually do not drink much water. Their bodies are
The physiological differences between these kidney types coated with mucus, which helps stem inward water movement.
are primarily related to the number of blood-filtering units they They absorb salts and ions by active transport across their gills.
contain. The pronephric kidney forms in the anterior portion of They also excrete a large volume of water as dilute urine.
the body cavity and contains fewer blood-filtering units than Although most groups of animals probably evolved in the
either the mesonephric or metanephric kidneys. The larger num- sea, many marine bony fishes probably had freshwater ances-
ber of filtering units in the latter has allowed vertebrates to face tors, as presented in chapter 18. Marine fishes face a different
the rigorous osmoregulatory and excretory demands of freshwater problem of water balance—their body fluids are hypoosmotic
and terrestrial environments. with respect to seawater (see table 28.2), and water tends to leave
What follows is a presentation of how a few vertebrates their bodies, resulting in dehydration (figure 28.16b). To com-
maintain their water and solute concentrations in different pensate, marine fishes drink large quantities of water, and they
habitats—in the seas, in freshwater, and on land (table 28.2). secrete Na, Cl, and Kions through secretory cells in their
(a)
FIGURE 28.15 (b)

Types of Kidneys in Vertebrates and Their Association with the Male
Reproductive System. The brown portions of the drawings represent
the mesoderm that forms both the kidneys and gonads. Notice that it
extends much of the length of the body during development. (a) The
primitive pronephric kidney is found in adult hagfishes and embryonic
fishes and amphibians. It is anterior in the body and contains segmental
renal tubules that lead from the body of the pronephros to the
archinephric duct. Notice that the testes are separated from the kidneys.
(b) The mesonephros is the functional kidney in the amniote embryo,
adult fishes, and amphibians. It is structurally similar to the nonseg-
mented opisthonephric (advanced mesonephric) kidney of most nonam-
niote vertebrates, such as sharks. The anterior portion of the
opisthonephros functions in blood cell formation and secretion of sex
hormones. Notice that the testes occupy the position of the anterior
opisthonephros, and the archinephric duct carries both sperm and urine.
(c) The metanephric kidney of adult amniotes (reptiles, birds, and mam-
mals) is the most advanced kidney. Notice the separate ureters (new
ducts) for carrying urine. The archinephric duct becomes the ductus def-
erens for carrying sperm. The kidney is more compact and located more (c)
caudally in the body.
TA B L E 2 8 . 2
HOW VARIOUS VERTEBRATES MAINTAIN WATER AND SALT BALANCE
ORGANISM ENVIRONMENTAL URINE MAJOR KEY ADAPTATION

CONCENTRATION CONCENTRATION NITROGENOUS
RELATIVE TO RELATIVE TO WASTE
BODY FLUIDS BLOOD
Freshwater fishes Hypoosmotic Hypoosmotic Ammonia Absorb ions through gills

Saltwater fishes Hyperosmotic Isoosmotic Ammonia Secrete ions through gills
Sharks Isoosmotic Isoosmotic Ammonia Secrete ions through rectal gland
Amphibians Hypoosmotic Very hypoosmotic Ammonia and urea Absorb ions through skin
Marine reptiles Hyperosmotic Isoosmotic Ammonia and urea Secrete ions through salt gland
Marine mammals Hyperosmotic Very hyperosmotic Urea Drink some water
Desert mammals No comparison Very hyperosmotic Urea Produce metabolic water
Marine birds No comparison Weakly hyperosmotic Uric acid Drink seawater and use salt glands
Terrestrial birds No comparison Weakly hypersmotic Uric acid Drink freshwater
Glomerulus
Salts
H 2O
Kidney
Salts (diffusion)
Salts (transport)
Hypoosmotic urine
Food only
(large volume)
Hypoosmotic freshwater
FIGURE 28.17
(a) Freshwater teleosts Water and Ion Uptake in an Amphibian. Water can enter this frog
(hypertonic blood) via food, through its highly permeable skin, or from the urinary bladder.
The skin also actively transports ions such as Na and Cl from the en-
vironment. The kidney forms a dilute urine by reabsorbing Na and
Salt Cl ions. Urine then flows into the urinary bladder, where most of the
remaining ions are reabsorbed.
H 2O
Seawater 2ⴙ
Naⴙ, Clⴚ, Mg SO42ⴚ
Kⴙ freshwater stream of their birth and enter the sea. Instead of con-
Kidney tinuing to pump ions in, as they have done in freshwater, the
H2O and salmon must now rid their bodies of salt. Years later, these same
salt
salmon migrate from the sea to their freshwater home to spawn.
Food Isoosmotic urine
As they do, the pumping mechanisms reverse themselves.
(small volume)
Mg2ⴙ, SO42ⴚ, Ca2ⴙ, PO43ⴚ
Amphibians
Hyperosmotic seawater The amphibian kidney is identical to that of freshwater fishes (fig-
(b) Marine teleosts
ure 28.16a), which is not surprising, because amphibians spend a
(hypotonic blood) large portion of their time in freshwater, and when on land, they
tend to seek out moist places. Amphibians take up water and ions
FIGURE 28.16 in their food and drink, through the skin that is in contact with
Osmoregulation. Osmoregulation by (a) freshwater and (b) marine moist substrates, and through the urinary bladder (figure 28.17).
fishes. Large black arrows indicate passive uptake or loss of water or ions. This uptake counteracts what is lost through evaporation and pre-
Small black and white arrows indicate active transport processes at gill vents osmotic imbalance (see table 28.2).
membranes and kidney tubules. Insets of kidney nephrons depict adapta-
tions within the kidney. Water, ions, and small organic molecules are fil- The urinary bladder of a frog, toad, or salamander is an
tered from the blood at the glomerulus of the nephron. Essential important water and ion reservoir. For example, when the envi-
components of the filtrate can be reabsorbed within the tubule system of ronment becomes dry, the bladder enlarges for storing more urine.
the nephron. Marine fishes conserve water by reducing the size of the If an amphibian becomes dehydrated, a brain hormone causes
glomerulus of the nephron, and thus reducing the quantity of water and water to leave the bladder and enter the body fluid.
ions filtered from the blood. Ions can be secreted from the blood into the
kidney tubules. Marine fishes can produce urine that is isoosmotic with the
blood. Freshwater fishes have enlarged glomeruli and short tubule systems. Reptiles, Birds, and Mammals
They filter large quantities of water from the blood, and tubules reabsorb
some ions from the filtrate. Freshwater fishes produce a hypoosmotic urine. Reptiles, birds, and mammals all possess metanephric kidneys
(see figure 28.15c). Their kidneys are by far the most complex
animal kidneys, well suited for these animals’ high rates of
metabolism.
gills. Channels in plasma membranes of their kidneys actively In most reptiles, birds, and mammals, the kidneys can
transport the multivalent ions that are abundant in seawater remove far more water than can those in amphibians, and the
(e.g., Ca2, Mg2, SO42, and PO43) out of the extracellular kidneys are the primary regulatory organs for controlling the
fluid and into the nephron tubes. The ions are then excreted in a osmotic balance of the body fluids. Some desert and marine rep-
concentrated urine. tiles and birds build up high salt (NaCl) concentrations in their
Some fishes encounter both fresh- and saltwater during bodies because they consume salty foods or seawater, and they
their lives. Newborn Atlantic salmon swim downstream from the lose water through evaporation and in their urine and feces. To
Afferent arteriole
Nephron
Interlobular artery
and vein
Cortex
Medulla
Arcuate artery
and vein Interlobar artery
and vein
Renal artery
Renal pelvis
Renal vein
Ureter
(a)
Proximal convoluted
tubule
Glomerulus
Glomerular
capsule
Efferent arteriole Afferent arteriole
FIGURE 28.18
Water Retention by Countercurrent Heating and Cooling in a Mam- (b)
mal. (a) When this animal inhales, the cool, dry air passing through
its nose is heated and humidified. At the same time, its nasal tissues are FIGURE 28.19
cooled. (b) When the animal exhales, it gives up heat to the previously Filtration Device of the Metanephric Kidney. (a) Interior of a kidney,
cooled nasal tissue. The air carries less water vapor, and condensation showing the positioning of the nephron and the blood supply to and
occurs in the animal’s nose. from the kidney. (b) Glomerular capsule. Red arrows show that high
blood pressure forces water and ions through small perforations in the
walls of the glomerular capillaries to form the glomerular filtrate.
rid themselves of excess salt, these animals also have salt glands
near the eye or in the tongue that remove excess salt from the nasal surfaces and does not leave the body. This mechanism
blood and secrete it as tearlike droplets. explains why a dog’s nose is usually cold and moist.
A major site of water loss in mammals is the lungs. To
reduce this evaporative loss, many mammals have nasal cavities
that act as countercurrent exchange systems (figure 28.18). When HOW THE METANEPHRIC KIDNEY
the animal inhales, air passes through the nasal cavities and is FUNCTIONS
warmed by the surrounding tissues. In the process, the tempera-
ture of this tissue drops. When the air gets deep into the lungs, it The filtration device of the metanephric kidney consists of over
is further warmed and humidified. During exhalation, as the warm one million individual filtration, secretion, and absorption struc-
moist air passes up the respiratory tree, it gives up its heat to the tures called nephrons (Gr. nephros, kidney on, neuter) (figure
nasal cavity. As the air cools, much of the water condenses on the 28.19a). At the beginning of the nephron is the filtration apparatus
FIGURE 28.20
Metanephric Nephron. The proximal convoluted tubule reabsorbs glucose and some ions. The distal convoluted tubule reabsorbs other ions and wa-
ter. Final water reabsorption takes place in the collecting duct. Black arrows indicate the direction of movement of materials in the nephron.
called the glomerular capsule (formerly Bowman’s capsule), which (ATP-requiring) and passive procedures are involved in the
looks rather like a tennis ball that has been punched in on one recovery of these substances. Potentially harmful compounds,
side (figure 28.19b). The capsules are in the cortical (outermost) such as hydrogen (H) and ammonium (NH4) ions, drugs, and
region of the kidney. In each capsule, an afferent (“going to”) arte- various other foreign materials are secreted into the nephron
riole enters and branches into a fine network of capillaries called lumen. In the last portion of the nephron, called the collecting
the glomerulus. The walls of these glomerular capillaries contain duct, final water reabsorption takes place so that the urine con-
small perforations called filtration slits that act as filters. Blood tains an ion concentration well above that of the blood. Thus, the
pressure forces fluid through these filters. The fluid is now known filtration, secretion, and reabsorption activities of the nephron do
as glomerular filtrate and contains small molecules, such as glu- not simply remove wastes. They also maintain water and ion bal-
cose, ions (Ca2, PO43), and the primary nitrogenous waste ance, and therein lies the importance of the homeostatic function
product of metabolism—urea or uric acid. Because the filtration of the kidney.
slits are so small, large proteins and blood cells remain in the Mammalian, and to a lesser extent avian and reptilian, kid-
blood and leave the glomerulus via the efferent (“outgoing”) arte- neys can remove far more water from the glomerular filtrate than
riole. The efferent arteriole then divides into a set of capillaries can the kidneys of amphibians. For example, human urine is four
called the peritubular capillaries that wind profusely around the times as concentrated as blood plasma, a camel’s urine is eight
tubular portions of the nephron (figure 28.20). Eventually, they times as concentrated, a gerbil’s is 14 times as concentrated, and
merge to form veins that carry blood out of the kidney. some desert rats and mice have urine more than 20 times as con-
Beyond the glomerular capsule are the proximal convoluted centrated as their plasma. This concentrated waste enables them
tubule, the loop of the nephron (formerly the loop of Henle), and to live in dry or desert environments, where little water is avail-
the distal convoluted tubule. At various places along these struc- able for them to drink. Most of their water is metabolically pro-
tures, the glomerular filtrate is selectively reabsorbed, returning duced from the oxidation of carbohydrates, fats, and proteins in
certain ions (e.g., Na, K, Cl) to the bloodstream. Both active the seeds that they eat (see table 28.1). Mammals and, to a lesser
extent, birds achieve this remarkable degree of water conser-

vation by a unique, yet simple, evolutionary adaptation: the
bending of the nephron tube into a loop. By bending, the
nephron can greatly increase the salt concentration in the tissue
through which the loop passes and use this gradient to draw
large amounts of water out of the tube.
Countercurrent Exchange
The loop of the nephron increases the efficiency of reabsorption
by a countercurrent flow similar to that in the gills of fishes or in
the legs of birds, but with water and ions being reabsorbed instead
of oxygen or heat. Generally, the longer the loop of the nephron,
the more water and ions that can be reabsorbed. It follows that
desert rodents (e.g., the kangaroo rat) that form highly concen-
trated urine have very long nephron loops (figure 28.21). Simi-
larly, amphibians that are closely associated with aquatic habitats
have nephrons that lack a loop.
FIGURE 28.21 Figure 28.22 shows the countercurrent flow mechanism for
Kangaroo Rat (Dipodomys ordii), a Master of Water Conservation. concentrating urine. The process of reabsorption in the proximal
Its efficient kidneys can concentrate urine 20 times that of its blood convoluted tubule removes some salt (NaCl) and water from the
plasma. As a result, these kidneys, as well as other adaptations, prevent
unnecessary water loss to the environment. glomerular filtrate and reduces its volume by approximately 25%.
Afferent Glomerular
arteriole capsule
H 2O H2O
300
Glomerulus Distal
Proximal – convoluted
Cl tubule H2 O
convoluted
Efferent +
tubule Na
arteriole – H 2O Cortex
Cl
+ Medulla
+ H 2O Na
Na active transport –
+ Cl
Na diffusion H2 O
Milliosmolar
+
– Na
Cl diffusion –
600 Cl
H2O diffusion H 2O Na
+ H 2O
–
Cl
H 2O Na
+ Collecting duct
–
Cl
+ H 2O
Na
– Urea
H 2O Cl Urea-brine bath
+
Na
Loop of the nephron Urea To urinary
bladder
1200
Freely permeable Impermeable Variably permeable

to water to water to water
FIGURE 28.22
Countercurrent Exchange. Movement of materials in the nephron and collecting duct. Solid arrows indicate active transport; dashed arrows indicate
passive transport. The shading at intervals along the tubules illustrates the relative concentration of the filtrate in milliosmoles.
However, the concentrations of salt and urea are still isosmotic

with the extracellular fluid.
As the filtrate moves to the descending limb of the loop of
the nephron, it becomes further reduced in volume and more con-
centrated. Water moves out of the tubule by osmosis due to the
high salt concentration (the “brine-bath”) in the extracellular
fluid.
Notice in figure 28.22 that the highest urea-brine bath con-
centration is around the lower portion of the loop of the nephron.
As the filtrate passes into the ascending limb, sodium (Na) ions
are actively transported out of the filtrate into the extracellular
fluid, with chloride (Cl) ions following passively. Water cannot
flow out of the ascending limb because the cells of the ascending
limb are impermeable to water. Thus, the salt concentration of the
extracellular fluid becomes very high. The salt flows passively into
the descending loop, only to move out again in the ascending
loop, creating a recycling of salt through the loop and the extra-
cellular fluid. Because the flows in the descending and ascending
limbs are in opposite directions, a countercurrent gradient in salt
is set up. The osmotic pressure of the extracellular brine bath is
made even higher because of the abundance of urea that moves
out of the collecting ducts.
Finally, the distal convoluted tubule empties into the col-
lecting duct, which is permeable to urea, and the concentrated FIGURE 28.23
urea in the filtrate diffuses out into the surrounding extracellular Component Parts of the Human Urinary System. The positions of
fluid. The high urea concentration in the extracellular fluid, cou- the kidneys, ureters, urinary bladder, and urethra.
pled with the high concentration of salt, forms the urea-brine
bath that causes water to move out of the filtrate by osmosis as it
moves down the descending limb. Finally, the many peritubular
capillaries surrounding each nephron collect the water and return opens at the body surface at the end of the penis (in human males)
it to the systemic circulation. or just in front of the vaginal entrance (in human females). As the
The renal pelvis of the mammalian kidney is continuous urinary bladder fills with urine, tension increases in its smooth
with a tube called the ureter that carries urine to a storage organ muscle walls. In response to this tension, a reflex response relaxes
called the urinary bladder (figure 28.23). Urine from two ureters sphincter muscles at the entrance to the urethra. This response is
(one from each kidney) accumulates in the urinary bladder. The called urination. The two kidneys, two ureters, urinary bladder,
urine leaves the body through a single tube, the urethra, which and urethra constitute the urinary system of mammals.
SUMMARY 6. The hypothalamus is the temperature regulating center that func-

tions as a thermostat with a fixed set point. This set point can ei-
1. Thermoregulation is a complex and important physiological ther rise or fall during hibernation or torpor.
process for maintaining heat homeostasis despite environmental 7. Some invertebrates have contractile vacuoles, flame-cell systems,
changes. antennal (green) glands, maxillary glands, coxal glands, nephridia,
2. Ectotherms generally obtain heat from the environment, whereas or Malpighian tubules for osmoregulation.
endotherms generate their own body heat from metabolic processes. 8. The osmoregulatory system of vertebrates governs the concentra-
3. Homeotherms generally have a relatively constant core body tem- tion of water and ions; the excretory system eliminates metabolic
perature, while heterotherms have a variable body temperature. wastes, water, and ions from the body.
4. The high, constant body temperature of birds and mammals also 9. Freshwater animals tend to lose ions and take in water. To avoid
depends on insulation, panting, sweating, specific behaviors, vaso- hydration, freshwater fishes rarely drink much water, have imper-
constriction or vasodilation of peripheral blood vessels, and in meable body surfaces covered with mucus, excrete a dilute urine,
some species, a rete mirabile system. and take up ions through their gills.
5. Thermogenesis involves mainly shivering, enzymatic activity, 10. Marine animals tend to take in ions from the seawater and to lose
brown fat, and high cellular metabolism. water. To avoid dehydration, they frequently drink water, have

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Miller−Harley: Zoology, III. Form and Function: A 27.

Nutrition and Digestion © The McGraw−Hill

NUTRITION AND DIGESTION

434 PART THREE Form and Function: A Comparative Perspective

EVOLUTION OF NUTRITION MACRONUTRIENTS

CHAPTER 27 Nutrition and Digestion 435

(a) (b) (c)

Carbohydrates Lipids Proteins

Glucose Steroids the Proteins the

Other lipids the Nitrogen

nerve and muscle in an animal’s body. Animals lose large

436 PART THREE Form and Function: A Comparative Perspective

in a special organ or cavity (ﬁgure 27.2b). Nutrients from the

CHAPTER 27 Nutrition and Digestion 437

VITAMIN CHARACTERISTICS FUNCTIONS SOURCES

438 PART THREE Form and Function: A Comparative Perspective

VITAMIN CHARACTERISTICS FUNCTIONS SOURCES

FIGURE 27.2 HERBIVORY

CHAPTER 27 Nutrition and Digestion 439

SURFACE NUTRIENT ABSORPTION

440 PART THREE Form and Function: A Comparative Perspective

Gastrovascular (c) Nematode

progressive digestive processing in specialized regions along

complete digestive tract correlate with different food-gathering Cytostome

ples further illustrate digestive systems in protozoa and 2. Excess water

CHAPTER 27 Nutrition and Digestion 441

442 PART THREE Form and Function: A Comparative Perspective

CHAPTER 27 Nutrition and Digestion 443

RUMENS In ruminants, the upper portion of the stomach expands to

444 PART THREE Form and Function: A Comparative Perspective

Lung duct Bile duct Spiral valve Intestine

Duct to swim bladder Gallbladder Stomach

Pharynx Pyloric ceca Intestine Anus

CHAPTER 27 Nutrition and Digestion 445

Rumen Esophagus Omasum Appendix

ments that enter the circulation. These pigments are subsequently

446 PART THREE Form and Function: A Comparative Perspective

Serosa Longitudinal Circular Mucosa

The process of digesting and absorbing nutrients in a mam-

CHAPTER 27 Nutrition and Digestion 447

448 PART THREE Form and Function: A Comparative Perspective

contraction that leads to peristalsis and segmentation, as well as

CHAPTER 27 Nutrition and Digestion 449

Esophagus Fundic region

Chief cell Submucosa

450 PART THREE Form and Function: A Comparative Perspective

CHAPTER 27 Nutrition and Digestion 451

452 PART THREE Form and Function: A Comparative Perspective

PLACE OF DIGESTION SOURCE SECRETION ENZYME DIGESTIVE FUNCTION

CHAPTER 27 Nutrition and Digestion 453

454 PART THREE Form and Function: A Comparative Perspective

ONLINE LEARNING CENTER • RELATED WEB LINKS

HOMEOSTASIS AND TEMPERATURE

456 PART THREE Form and Function: A Comparative Perspective

constant (homeostatic), relatively high body temperature is a

THE IMPACT OF TEMPERATURE Convection

Every animal’s physiological functions are inexorably linked to

CHAPTER 28 Temperature and Body Fluid Regulation 457

(have a variable body temperature), there are many exceptions.