Chapter 41

Animal Nutrition

Lecture Outline

Overview: The Need to Feed

 All animals eat other organisms—dead or alive, whole or by the piece (including parasites).
 Animal nutrition includes the ingestion, breakdown, and absorption of food.
 In general, animals fit into one of three dietary categories.
1. Herbivores, such as cattle, parrotfish, and termites, eat mainly plants or algae.
2. Carnivores, such as sharks, hawks, and spiders, eat other animals.
3. Omnivores, such as cockroaches, bears, and humans, consume animal and plant or algal
matter.
 The terms herbivore, carnivore, and omnivore represent the kinds of food that an animal usually eats,
but most animals are opportunistic, occasionally eating foods that are outside their main dietary
category.
o For example, cattle and deer, which are herbivores, occasionally eat small animals or bird
eggs.
o All animals consume bacteria along with other types of food.
 To survive and reproduce, animals must balance their consumption, storage, and use of food.

Concept 41.1 An animal’s diet must supply chemical energy, organic molecules, and
essential nutrients.
 An animal’s diet provides chemical energy that can be converted to ATP to power living
processes.
 In addition to fuel for ATP production, an animal’s diet must supply the raw materials needed
for biosynthesis.
o Animals require a source of organic carbon and a source of organic nitrogen in order to
construct their own organic molecules.
 Materials that an animal’s cells require but cannot synthesize are called essential nutrients.
o Essential nutrients, which must be obtained from an animal’s diet, include both minerals and
preassembled organic molecules.
o Some nutrients are essential for all animals, whereas others are required for only certain
species.
 Overall, an adequate diet must supply chemical energy for cellular processes, organic molecules
as building blocks for macromolecules, and essential nutrients.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-1
 There are four classes of essential nutrients: essential amino acids, essential fatty acids, vitamins,
and minerals.
 Essential amino acids are those an animal cannot synthesize.
○ Most animals require eight amino acids in their diet.
 A diet that does not provide sufficient amounts of one or more essential amino acids leads to
protein deficiency, the most common type of malnutrition.
 Animal proteins are “complete,” providing all the essential amino acids in their proper
proportions.
 Most plant proteins are “incomplete,” deficient in one or more essential amino acids.
 Animals can synthesize most of the fatty acids they need.
 Essential fatty acids, the ones that animals cannot synthesize, are unsaturated.
 Vitamins are organic molecules with diverse functions that are required in the diet in quantities
that are quite small compared with the relatively large quantities of essential amino acids and
fatty acids that animals need.
o Although vitamins are required in tiny amounts—from about 0.01 mg to 100 mg per day—
depending on the vitamin, vitamin deficiency (or overdose in some cases) can cause serious
problems.
○ So far, 13 vitamins essential to humans have been identified.
 Vitamins can be grouped into water-soluble vitamins and fat-soluble vitamins, with extremely
diverse physiological functions.
 One water-soluble vitamin is the B complex, which consists of several compounds that function
as coenzymes in key metabolic processes.
 Vitamin C, also water-soluble, is required for the production of connective tissue.
 Excessive amounts of water-soluble vitamins are excreted in urine, and moderate overdoses are
probably harmless.
 The fat-soluble vitamins are A, D, E, and K. They have a wide variety of functions.
○ Vitamin A is incorporated in the visual pigments of the eye.
○ Vitamin D aids in calcium absorption and bone formation.
○ Vitamin E seems to protect membrane phospholipids from oxidation.
○ Vitamin K is required for blood clotting.
 Excess amounts of fat-soluble vitamins are not excreted but are deposited in body fat.
o Overconsumption may lead to toxic accumulations of these compounds.
 The subject of vitamin dosage has aroused heated scientific and popular debate.
 Some believe that it is sufficient to meet the recommended daily allowances (RDAs), the nutrient
intake proposed by nutritionists to maintain health.
 Others argue that the RDAs are set too low for some vitamins, and a fraction of these people
believe, probably mistakenly, that massive doses of vitamins confer health benefits.
 Debate centers on the optimal doses of vitamins C and E.
 Research is ongoing, but all that can be said with any certainty is that people who eat a balanced
diet are not likely to develop symptoms of vitamin deficiency.
 Minerals are simple inorganic nutrients, usually required in small amounts—from less than 1 mg
to about 2,500 mg per day.
 Mineral requirements vary with animal species.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-2
 Humans and other vertebrates require relatively large quantities of calcium and phosphorus for
the construction and maintenance of bone.
o Calcium is also necessary for the normal functioning of nerves and muscles.
o Phosphorus is a component of the cytochromes that function in cellular respiration.
 Iron is a component of the cytochromes that function in cellular respiration and of hemoglobin,
the oxygen-binding protein of red blood cells.
 Magnesium, iron, zinc, copper, manganese, selenium, and molybdenum are cofactors built into
the structure of certain enzymes.
○ Magnesium, for example, is present in enzymes that split ATP.
 Iodine is required for thyroid hormones, which regulate metabolic rate.
 Sodium, potassium, and chloride are important in nerve function and have a major influence on
the osmotic balance between cells and interstitial fluids.
 Excess consumption of some minerals can upset the homeostatic balance and cause toxic effects.
o Liver damage due to iron accumulation affects up to 10% of the population in regions of
Africa that have an iron-rich water supply.
o Excess consumption of salt (sodium chloride) may contribute to high blood pressure.
o The average U.S. citizen eats enough salt to provide about 20 times the required amount of
sodium.
 When an animal is undernourished, it uses up stored fat and carbohydrates, and the body begins
breaking down its own proteins for fuel.
o Muscles begin to decrease in size, and the brain can become protein-deficient.
o If energy intake remains less than energy expenditure, death will eventually result.
o Even if a seriously undernourished person survives, some damage may be irreversible.
 Because a diet of a single staple such as rice or corn can often provide sufficient calories,
undernourishment is generally common when drought, war, or some other crisis severely
disrupts the food supply.
 Another cause of undernourishment is anorexia nervosa, an eating disorder associated with
compulsive starvation despite food availability.
 The potential effects of malnourishment include deformities, disease, and even death.
o For example, cattle, deer, and other herbivores may develop fragile bones if they graze in
areas where soils and plants are deficient in phosphorus.
o Some grazing animals obtain the missing nutrients by consuming concentrated sources of
salt or other minerals.
 Recent experiments with spiders have found that carnivores can adjust for dietary deficiencies by
switching to prey that restores their nutritional balance.
 Humans can suffer from malnourishment.
o People subsisting on a rice diet may suffer from vitamin A deficiency, which can cause
blindness or death.
o A genetically engineered strain of “Golden Rice” synthesizes beta-carotene, a source of
vitamin A.
 It is difficult to determine an ideal human diet. Humans are genetically diverse and live in varied
settings.
 Ethical concerns preclude experimenting on the nutritional needs of children.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-3
 Researchers study genetic defects that disrupt food uptake, storage, or use.
o Hemochromatosis causes a buildup of iron in the absence of excess iron consumption.
o By studying this disease, scientists have learned about the genes that regulate iron
absorption.
 Insights into human nutrition have come from epidemiology, the study of human health and disease
at the population level.
o Epidemiologists aim to identify nutritional strategies for the prevention and control of
diseases and disorders.
 In the 1970s, researchers found that women of low socioeconomic status are more likely to have
children with neural tube defects, which occur when tissue fails to enclose the developing brain
and spinal cord.
 Richard Smithells of the University of Leeds found that folic acid (vitamin B 9 ) supplements
greatly reduce the risk of neural tube defects.
 In 1998, the FDA began to require that folic acid be added to the enriched grain used to make
bread and cereal.
 The frequency of neural tube defects has been significantly reduced as a result.

Concept 41.2 The main stages of food processing are ingestion, digestion,
absorption, and elimination.
 Food processing by animals can be divided into four distinct stages: ingestion, digestion,
absorption, and elimination.
 Ingestion, the act of eating, is the first stage of food processing.
 Food can be ingested in many liquid and solid forms.
 Digestion, the second stage of food processing, is the process of breaking food down into
molecules small enough for the body to absorb.
o Food must be broken down because animals cannot directly use the macromolecules in
food. They are too large to pass through the cell membranes to enter the cells of the animal.
o In addition, the proteins, carbohydrates, nucleic acids, fats, and phospholipids in food are
not identical to those an animal makes itself.
 Digestion cleaves macromolecules into their component monomers, which the animal then uses
to make its own molecules or as fuel for ATP production.
 Digestion reverses the process that a cell uses to link together monomers to form
macromolecules.
o Rather than removing a molecule of water for each new covalent bond formed, digestion
breaks bonds with the addition of water via enzymatic hydrolysis.
 A variety of hydrolytic enzymes catalyze the digestion of each of the classes of macromolecules
found in food.
o Polysaccharides and disaccharides are split into simple sugars.
o Fats are digested to glycerol and fatty acids.
o Proteins are broken down into amino acids.
o Nucleic acids are cleaved into nucleotides.
 Chemical digestion is usually preceded by mechanical fragmentation of the food—by chewing,
for instance.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-4
 Breaking food into smaller pieces increases the surface area exposed to digestive juices
containing hydrolytic enzymes.
 After the food is digested, the animal’s cells take up small molecules such as amino acids and
simple sugars from the digestive compartment, a process called absorption.
 During elimination, undigested material passes out of the digestive compartment.
 To avoid digesting their own cells and tissues, most organisms carry out digestion in specialized
compartments.
 The simplest digestive compartments are food vacuoles, organelles in which hydrolytic enzymes
break down food without digesting the cell’s own cytoplasm, a process termed intracellular
digestion.
o This process begins after a cell has engulfed food by phagocytosis or pinocytosis.
 Newly formed food vacuoles fuse with lysosomes, which are organelles containing hydrolytic
enzymes.
 A few animals, such as sponges, digest their food entirely by this mechanism.
 In most animals, at least some hydrolysis occurs by extracellular digestion, the breakdown of
food outside cells.
o Extracellular digestion occurs within compartments that are continuous with the outside of
the animal’s body.
o Thus, organisms can devour much larger prey than can be ingested by phagocytosis.
 Many animals with simple body plans have digestive sacs with single openings, called
gastrovascular cavities.
 Gastrovascular cavities function in both the digestion and distribution of nutrients throughout
the body.
o For example, the cnidarians called hydras capture their prey with nematocysts and use
tentacles to stuff the prey through the mouth into the gastrovascular cavity.
o The prey is then partially digested by enzymes secreted by specialized gland cells of the
gastrodermis.
o Nutritive muscular cells in the gastrodermis engulf the food particles.
o Most of the actual hydrolysis of macromolecules occurs intracellularly.
o Undigested materials are eliminated through the mouth.
 In contrast to cnidarians and flatworms, most animals have digestive tubes extending between a
mouth and anus.
 These digestive tubes are called complete digestive tracts, or alimentary canals.
 Because food moves in one direction, the tube can be organized into specialized regions that
carry out digestion and nutrient absorption in a stepwise fashion.
 In addition, animals with alimentary canals can eat more food before the earlier meal is
completely digested.

Concept 41.3 Organs specialized for successive stages of food processing form the
mammalian digestive system.
 The general principles of food processing are similar for a diversity of animals, including
mammals. We will use the mammalian system as a representative example.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-5
 The mammalian digestive system consists of the alimentary canal and various accessory glands
that secrete digestive juices into the canal through ducts.
o The accessory glands include the salivary glands, the pancreas, the liver, and the gallbladder.
 Peristalsis, rhythmic waves of contraction by smooth muscles in the walls of the canal, pushes
food along.
 Sphincters, muscular ring-like valves, regulate the passage of material between specialized
chambers of the canal.
The oral cavity, pharynx, and esophagus initiate food processing.
 Both physical and chemical digestion of food begins in the mouth, or oral cavity.
 During chewing, teeth of various shapes cut, smash, and grind food, making it easier to swallow
and increasing its surface area.
 The presence of food in the oral cavity triggers a nervous reflex that causes the salivary glands
to deliver saliva through ducts to the oral cavity.
o Salivation may occur in anticipation of food because of learned associations between eating
and the time of day, cooking odors, or other stimuli.
 Chemical digestion of carbohydrates, a main source of chemical energy, begins in the oral cavity.
 Saliva contains amylase, an enzyme that hydrolyzes starch and glycogen into smaller
polysaccharides and the disaccharide maltose.
 Saliva contains a slippery glycoprotein called mucin, which protects the soft lining of the mouth
from abrasion and lubricates the food for easier swallowing.
 Saliva also contains buffers that help prevent tooth decay by neutralizing acid in the mouth.
 Antibacterial agents in saliva, such as lysozyme, kill microbes that enter the mouth with food.
 The tongue tastes food, manipulates it during chewing, and helps shape the food into a ball
called a bolus.
 During swallowing, the tongue pushes a bolus back into the oral cavity and into the pharynx.
 The pharynx, also called the throat, is a junction that opens to both the esophagus and the
trachea (windpipe).
 When we swallow, the top of the windpipe moves up so that its opening, the glottis, is blocked by
a cartilaginous flap, the epiglottis.
 This mechanism normally ensures that a bolus will be guided into the entrance of the esophagus
and not directed down the windpipe.
○ If food or liquid enters and blocks the windpipe, the material can be dislodged by vigorous
coughing or a forced upward thrust of the diaphragm (the Heimlich maneuver).
 The esophagus contains both striated and smooth muscle.
o The striated muscle at the top of the esophagus is active during swallowing.
o In the rest of the esophagus, smooth muscle functions in peristalsis, as rhythmic cycles of
contraction move the bolus to the stomach.
The stomach stores food and performs preliminary digestion.
 The stomach is in the upper abdominal cavity, just below the diaphragm.
 With accordion-like folds and a very elastic wall, the stomach can stretch to accommodate about
2 L of food and fluid, storing an entire meal.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-6
 The stomach secretes a digestive fluid called gastric juice and mixes this secretion with the food
by the churning action of the smooth muscles in the stomach wall.
o The mixture of ingested food and digestive juices is called chyme.
 Two components of gastric juice carry out chemical digestion in the stomach.
 One component of gastric juice is hydrochloric acid (HCl), which disrupts the extracellular
matrix that binds cells together.
o With a high concentration of hydrochloric acid, the gastric juice has a pH of about 2—acidic
enough to digest iron nails.
o This low pH kills most bacteria that are swallowed with food.
o It also denatures proteins in food, increasing exposure of their peptide bonds.
 The second component of gastric juice is pepsin, an enzyme that begins the hydrolysis of
proteins.
o Pepsin, which works well in strongly acidic environments, is a protease that breaks peptide
bonds adjacent to specific amino acids, producing smaller polypeptides.
 Cells in the gastric glands of the stomach produce the components of gastric juice.
o Parietal cells secrete hydrochloric acid in the form of hydrogen and chloride ions, using an
ATP-driven pump.
o Meanwhile, chief cells release pepsin into the lumen in an inactive form called pepsinogen.
○ HCl in the lumen of the stomach converts pepsinogen to active pepsin by clipping off a
small portion of the molecule to expose its active site.
○ In a positive-feedback system, activated pepsin activates more pepsinogen molecules.
 Because HCl and pepsin form in the lumen of the stomach, not within the cells of the gastric
glands, the stomach’s cells are protected from self-digestion.
 The stomach’s second line of defense against self-digestion is a coating of mucus, secreted by
the epithelial cells, that protects the stomach lining.
o Still, the epithelium is continuously eroded, and the epithelium is completely replaced by
mitosis every three days.
 Gastric ulcers, lesions in the stomach lining, are caused by the acid-tolerant bacterium Helicobacter
pylori.
o Ulcers are often treated with antibiotics.
o The discovery that ulcers are caused by a bacterial infection, not by stress, earned Barry
Marshall and J. Robin Warren the Nobel Prize in 2005.
 About every 20 seconds, the stomach contents are mixed by the churning action of smooth
muscles.
 As a result of mixing and enzyme action, what begins in the stomach as a recently swallowed
meal becomes nutrient-rich chyme.
 Most of the time, the stomach is closed off at both ends.
 The opening from the esophagus to the stomach normally dilates only when a bolus driven by
peristalsis arrives.
o The occasional backflow of acid chyme from the stomach into the lower esophagus—
known as acid reflux—causes the irritation of the esophagus called “heartburn.”
 The sphincter at the opening from the stomach to the small intestine helps regulate the passage
of chyme into the intestine.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-7
 o A squirt at a time, it takes about 2 to 6 hours after a meal for the stomach to empty.
The small intestine is the major organ of digestion and absorption.
 With a length of more than 6 m in humans, the small intestine is the longest section of the
alimentary canal.
 Most of the enzymatic hydrolysis of food macromolecules and most of the absorption of
nutrients into the blood occur in the small intestine.
 In the duodenum, the first 25 cm or so of the small intestine, chyme from the stomach mixes
with digestive juices from the pancreas, liver, gallbladder, and gland cells of the intestinal wall.
 Hormones released by the stomach and duodenum control the digestive secretions into the
alimentary canal.
 The pancreas produces several enzymes and an alkaline solution rich in bicarbonate that acts as
a buffer to neutralize the acidity of chyme from the stomach.
o Pancreatic enzymes include trypsin and chymotrypsin, proteases that are secreted into the
duodenum in inactive form.
o The pancreatic proteases are activated once they are in the extracellular space within the
duodenum.
 The liver performs a wide variety of important functions in the body, including the production
of bile.
o Bile contains bile salts that act as detergents that aid in the digestion and absorption of fats.
o Bile is stored and concentrated in the gallbladder.
 The liver also breaks down toxins that enter the body and helps balance nutrient utilization.
○ Bile also contains pigments that are by-products of red blood cell destruction in the liver.
○ These bile pigments are eliminated from the body with the feces.
 The brush border of the epithelial lining of the duodenum produces several digestive enzymes.
o Some enzymes are secreted into the lumen, while others are bound to the surface of the
epithelial cells.
 Enzymatic digestion is completed as peristalsis moves the mixture of chyme and digestive juices
along the small intestine.
 Most digestion is completed while the chyme is still in the duodenum.
 The remaining regions of the small intestine, the jejunum and ileum, function mainly in the
absorption of nutrients and water.
 To enter the body, nutrients in the lumen must pass the lining of the digestive tract.
 A few nutrients are absorbed in the stomach and large intestine, but most absorption takes place
in the small intestine.
 The enormous surface area of the small intestine is an adaptation that greatly increases the rate
of nutrient absorption.
o The small intestine has a surface area of 300 m2, roughly the size of a tennis court.
 Large circular folds in the lining bear fingerlike projections called villi.
 Each epithelial cell of a villus has many microscopic appendages called microvilli that are
exposed to the intestinal lumen.
o The microvilli are the basis of the term brush border for the intestinal epithelium.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-8
 In some cases, transport of nutrients across the epithelial cells is passive, as molecules move
down their concentration gradients from the lumen of the intestine into the epithelial cells, and
then into capillaries.
o Fructose, a simple sugar, moves by facilitated diffusion down its concentration gradient from
the lumen of the intestine into the epithelial cells and then into capillaries.
 Amino acids, small peptides, vitamins, and most glucose molecules are pumped against
concentration gradients by the epithelial cells of the villus.
o This active transport allows much more absorption of nutrients than would be possible with
passive diffusion.
 Monoglycerides and most fatty acids are absorbed by epithelial cells and recombined into
triglycerides within the cells.
 The fats are mixed with cholesterol and coated with special proteins to form small globules
called chylomicrons.
 Chylomicrons are transported by exocytosis out of epithelial cells and into lacteals, lymph
vessels at the core of each villus.
 The lacteals converge into the larger vessels of the lymphatic system, eventually draining into
large veins that return blood to the heart.
 The capillaries and veins that drain nutrients away from the villi converge into the hepatic
portal vein, which leads directly to the liver.
 Therefore, the liver, which has the metabolic versatility to interconvert various organic
molecules, has first access to the amino acids and sugars absorbed after a meal is digested.
 The liver modifies and regulates this varied mix before releasing materials back into the
bloodstream.
o The liver helps regulate the levels of glucose in the blood, ensuring that the blood exiting the
liver usually has a glucose concentration very close to 90 mg per 100 mL, regardless of the
carbohydrate content of the meal.
o The liver also removes toxic substances before the blood circulates broadly.
 From the liver, blood travels to the heart, which pumps the blood and nutrients to all parts of
the body.
Absorption of water is the major function of the large intestine.
 The large intestine, or colon, is connected to the small intestine at a T-shaped junction where a
sphincter controls the movement of materials.
 One arm of the T is a pouch called the cecum.
o The relatively small cecum of humans has a fingerlike extension, the appendix, which makes
a minor contribution to body defense.
 The main branch of the human colon is shaped like an upside-down U, about 1.5 m long.
 A major function of the colon is to recover water that has entered the alimentary canal as the
solvent in various digestive juices.
o About 7 L of fluid are secreted into the lumen of the digestive tract of a person each day.
o More than 90% of the water is reabsorbed in the small intestine and the colon.
 Because there is no biological mechanism for the active transport of water, water absorption in
the colon occurs by osmotically driven bulk flow as ions, particularly sodium, are pumped out of
the lumen.
 Digestive wastes, the feces, become more solid as they are moved along the colon by peristalsis.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-9
 o It takes 12 to 24 hours for material to travel the length of the organ.
o If the lining of the colon is irritated by a viral or bacterial infection, less water than usual is
resorbed, resulting in diarrhea.
o If too much water is absorbed because peristalsis moves the feces too slowly, the result is
constipation.
 Living in the large intestine is a rich flora of mostly harmless bacteria, which make up
approximately one-third of the dry weight of feces.
o One of the most common inhabitants of the human colon is Escherichia coli, a favorite
research organism.
○ Because E. coli is so common in human digestive systems, its presence in lakes and streams is
an indicator of contamination by untreated sewage.
 As a by-product of their metabolism, many colon bacteria generate gases, including methane and
hydrogen sulfide.
 These gases and ingested air are expelled through the anus.
 Some bacteria produce vitamins, including biotin, folic acid, vitamin K, and several B vitamins,
which supplement our dietary intake of vitamins.
 Feces contain masses of bacteria and undigested materials, including cellulose.
o Although cellulose fibers have no caloric value to humans, their presence in the diet helps
move food along the digestive tract.
 The terminal portion of the colon is called the rectum, where feces are stored until they can be
eliminated.
 Between the rectum and the anus are two sphincters, one involuntary and one voluntary.
 Once or more each day, strong contractions of the colon create an urge to defecate.

Concept 41.4 Evolutionary adaptations of vertebrate digestive systems correlate with

diet.
The variety of digestive systems correlates with diet.
 The digestive systems of mammals and other vertebrates are variations on a common plan.
 The many intriguing varieties of digestive systems are often associated with the animal’s diet.
 Dentition, an animal’s assortment of teeth, is one example of structural variation reflecting diet.
 Mammals’ evolutionary adaptation of teeth for processing different kinds of food is one of the
major reasons they have been so successful.
 Nonmammalian vertebrates generally have less specialized dentition, but there are exceptions.
o For example, poisonous snakes, such as rattlesnakes, have fangs, modified teeth that inject
venom into prey.
o Some snakes have hollow fangs, like syringes, while others drip poison along grooves in the
tooth surface.
 All snakes have another important anatomical adaptation for feeding, the ability to swallow large
prey whole.
o The lower jaw of a snake is loosely hinged to the skull by an elastic ligament that permits the
mouth and throat to open very wide for swallowing.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-10
 Large, expandable stomachs are common in carnivores, which may go for a long time between
meals and, therefore, must eat as much as they can when they do catch prey.
o For example, a 200-kg African lion can consume 40 kg of meat in one meal.
 The length of the vertebrate digestive system is also correlated with diet.
 In general, herbivores and omnivores have longer alimentary canals relative to their body sizes
than do carnivores, providing more time for digestion and more surface areas for absorption of
nutrients.
o Vegetation is more difficult to digest than meat because it contains cell walls. A longer
digestive tract allows more time for digestion and more surface area for absorption.
Symbiotic microorganisms help nourish many vertebrates.
 Much of the chemical energy in the diet of herbivorous animals is contained in the cellulose of
plant cell walls.
 Animals do not produce enzymes that hydrolyze cellulose, however, so many vertebrates (and
termites) house large populations of symbiotic bacteria and protists in special fermentation
chambers in their alimentary canals.
o These microbes have enzymes that can digest cellulose to simple sugars that the animal can
absorb.
 The location of symbiotic microbes in herbivores’ digestive tracts varies depending on the
species.
o The hoatzin, an herbivorous bird that lives in South American rain forests, has a large,
muscular crop (an esophageal pouch) that houses symbiotic microbes.
o Many herbivorous mammals, including horses, house symbiotic microbes in a large cecum,
the pouch where the small and large intestines connect.
o The symbiotic bacteria of rabbits and some rodents live in the large intestine and cecum.
 Since most nutrients are absorbed in the small intestine, these herbivores recover nutrients from
fermentation in the large intestine by coprophagy, eating some of their feces and passing food
through the alimentary canal a second time.
o The koala also has an enlarged cecum, where symbiotic bacteria ferment finely shredded
eucalyptus leaves.
 The most elaborate adaptations for an herbivorous diet have evolved in the ruminants, which
include deer, cattle, and sheep.
Adaptations related to digestion are widespread among multicellular animals.
 The giant tubeworms that live at deep-sea hydrothermal vents thrive at pressures of 260
atmospheres and temperatures of 4000C.
 These worms have no mouth or digestive systems. Instead, they rely on symbiotic bacteria to
generate energy and nutrients from the carbon dioxide, oxygen, hydrogen sulfide, and nitrate
available at the vents.
 For invertebrates and vertebrates alike, symbiosis has evolved as a general strategy for expanding
the sources of nutrition available for animals.

Concept 41.5 Homeostatic mechanisms contribute to an animal’s energy balance.

 Energy from food balances the expenditure of energy for metabolism, activity, and storage.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-11
 Nearly all ATP generation is based on the oxidation of organic fuel molecules—carbohydrates,
proteins, and fats—in cellular respiration.
 The monomers of any of these substances can be used as fuel, but most animals “burn” proteins
only after exhausting their supply of carbohydrates and fats.
 Fats are especially rich in energy, liberating about twice the energy liberated from an equal
amount of carbohydrate or protein during oxidation.
 When animals take in more energy-rich molecules than they break down, the excess is converted
into storage molecules such as glycogen.
 In humans, glycogen is stored in liver and muscle cells.
o Glycogen synthesis and breakdown are tightly regulated by the hormones insulin and
glucagon.
 Adipose cells are a secondary site of energy storage in the body.
 If glycogen stores are full and caloric intake still exceeds caloric expenditure, the excess is usually
stored as fat.
 When fewer calories are taken in than are expended—perhaps because of sustained heavy
exercise or lack of food—fuel is taken out of storage depots and oxidized.
 The human body expends liver glycogen first and then draws on muscle glycogen and fat.
o Most healthy people—even if they are not obese—have enough stored fat to sustain them
through several weeks of starvation.
 Overnourishment, the consumption of more food energy than the body needs for normal
metabolism, causes obesity, the excessive accumulation of fat.
 Obesity contributes to a number of health problems, including type 2 diabetes, colon and breast
cancer, and cardiovascular disease.
o Obesity is a contributing factor in 300,000 deaths per year in the United States.
 Researchers have discovered several homeostatic mechanisms that help regulate body weight.
 Operating as feedback circuits, these mechansms control the storage and metabolism of fat.
 Several hormones regulate long-term and short-term appetite by affecting a “satiety center” in
the brain.
o Mice with mutations in the ob or db gene eat voraciously and become obese.
o Doug Coleman determined that the ob gene is needed to produce the satiety factor and the
db gene is required to respond to the factor.
○ Cloning of the ob gene has shown that it codes for the hormone leptin.
o The db gene encodes the leptin receptor.
 Adipose cells produce leptin. As the amount of adipose tissue increases, leptin production
increases.
 High leptin levels cue the brain to suppress appetite.
 Conversely, loss of body fat decreases leptin levels in the blood, signaling the brain to increase
appetite.
 It is far from clear that our understanding of leptin will lead to effective treatments for obesity.
o Leptin has complex functions and plays a role in the formation of the nervous system.
o In addition, very few obese people have defective leptin production.
o In fact, most obese humans have abnormally high leptin levels, due to their large amounts of
adipose tissue.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-12
 o For some reason, the brain’s satiety center does not respond to the high leptin levels in many
obese people.
 Most humans crave fatty foods. Although fat hoarding is a health liability today, it may have been
advantageous in our evolutionary past.
o Our ancestors on the African savanna were hunter-gatherers who probably survived mainly
on plant materials, occasionally supplemented by meat.
o Natural selection may have favored those individuals with a physiology that induced them to
gorge on fatty foods on the rare occasions they were available.
o Individuals with genes that promoted the storage of fat during feasts may have been more
likely to survive famine.
 Obesity may be beneficial in certain species.
 Small seabirds called petrels fly long distances to find food that is rich in lipids.
 By bringing lipid-rich food to their chicks, the parents minimize the weight of food that they
must carry.
 Because these foods are low in protein, however, young petrels have to consume more calories
than they burn in metabolism—and consequently they become obese.
 Their fat deposits may help them to survive periods when parents cannot find enough food.
 In some petrel species, chicks at the end of the growth period weigh much more their parents.
They are too heavy to fly and must first starve for several days.

Lecture Outline for Campbell/Reece Biology, 8th Edition, © Pearson Education, Inc. 41-13