1 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Chapter 40 Animal Metabolism, Nutrition, and

Digestion
The biochemical pathways that make up animal metabolism are remarkably conserved among diverse groups of organisms.

These pathways evolved early in the history of life.

Nearly all animals depend on three main molecules as sources of energy and building blocks for growth and development: carbohydrates, fats, and
proteins.

The conversion of these food sources into biologically useful forms typically involves synthesizing adenosine triphosphate (ATP).

The breakdown of carbohydrates, fats, and proteins to produce ATP involves a linked set of chemical reactions.

Reactions that break down food sources to fuel the energy needs of a cell are referred to as catabolic.

Reactions that result in net energy storage within cells and the organism are anabolic.

Cellular Respiration: digesting organic compounds to

release stored ATP.
Glucose can be partially broken down in the absence of oxygen by
glycolysis.

Occurs in the cytosol of the cell

Breakdown of each glucose molecule results in

two molecules of pyruvate and

two molecules of ATP

Anaerobic metabolism

If oxygen is not present, or is present only in small amounts,

pyruvate is converted by fermentation into lactic acid.

The production of ATP by anaerobic metabolism provides rapid but

short-term energy to the cell and organism.

Aerobic metabolism

Carried out within the mitochondria of eukaryotic animals

Requires oxygen diffusing into the mitochondria

Provides a steady supply of ATP for longer-term, sustainable

activity.

Pyruvate is processed by the citric acid cycle rather than converted to lactic acid.

The electron transport chain and oxidative phosphorylation result in ATP production.

Oxygen is consumed and water is produced.

Lipids are another important energy source for most animals. Lipids, such as triacylglycerol, consumed in the diet are broken down to glycerol and free
fatty acids, which enter glycolysis or the citric acid cycle to yield ATP by mitochondrial electron transport.

Proteins consumed in the diet are also a useful energy source for animals. Proteins, which are needed for building and maintaining the body, constitute
the enzymes and structural elements of cells and tissues. Only following prolonged food deprivation, when fat and carbohydrate reserves are depleted,
do animals break down protein reserves to form ATP. Animals preferentially rely on fats for long-term energy supply.

Metabolic Rate: speed of energy usage. Variable. Aerobic metabolism can be measured by
oxygen consumption.
An animals overall rate of energy use defines its metabolic rate.

Metabolic rate can be measured by the animals rate of oxygen consumption, which in turn reflects the aerobic production of ATP.

Metabolic rate is affected by many factors, one of which is the activity level of an organism.

When an animal shifts from rest to activity, its metabolic rate and oxygen consumption rise to meet its increased demand for ATP.

The onset of activity requires immediate energy, which in animals is provided by specialized energy stores in their tissues.

Although glycolysis produces relatively few ATP molecules per molecule of glucose broken down, the reactions are extremely fast, providing a rapid
short-term supply of energy for animals.
2 Chapter 40 Animal Metabolism, Nutrition, and Digestion
Animals rely on anaerobic glycolysis for short bursts of intensive activity.

O2 Consumption During Activity: most exercise ATP requirements are

met by aerobic respiration, but prolonged activity may rely on
anaerobic respiration to make ATP, and this results in lactic acid
accumulation.
Eventually, enough oxygen diffuses into the
mitochondria to allow aerobic respiration to occur at a
rate that meets the ATP needs of the animal.
Therefore, the rate of oxygen consumption initially increases, then levels off. At
this point, the animals need for energy is being met entirely by aerobic
respiration.

With even more prolonged or intense activity, more ATP is needed, and it is harder
to meet that need by aerobic metabolism alone. More intensive physical exercise
requires a greater reliance on anaerobic glycolysis to produce ATP. The resulting
buildup of lactic acid and decrease in pH inside the muscle cells force an animal to
decrease its activity. The production of lactic acid through fermentation limits an
animals performance by lowering the pH of the animals blood, producing a
condition called metabolic acidosis, which contributes to fatigue.

When activity ends, the animals oxygen consumption rate declines but does not
immediately return to resting levels, The elevated consumption of oxygen following activity is the animals recovery metabolism. It
represents the continued metabolic energy required to reestablish the resting metabolic state of
the cells. During recovery metabolism, cells re- synthesize depleted ATP stores and metabolize the
end products of fermentation, particularly lactic acid.

The difference between an animals immediate energy need at the onset of activity versus energy supplied by aerobic
metabolism is often referred to as the animals oxygen debt. This debt is paid back following exercise by the animals recovery
metabolism. Recovery metabolism is associated with elevated breathing and heart rates are experienced when resting after
moderate to intense exercise.

Body Temperature and Activity: second law of thermodynamics energy

conversion converts most of the energy to
heat.
Metabolic rate increases with activity level. A by-product of metabolism is the
generation of heat, which is a necessary consequence of the second law of
thermodynamics. Animals have several mechanisms for dissipating excess heat,
including sweating, changes in blood flow, and panting.

Cheetahs, as top predators, are one of the fastest land animals known today,
capable of speeds up to 70 mph (114 km/h). Such extreme speed generates a lot
of heat. Interestingly, instead of releasing the excess heat as it builds up, the cheetah stores it, dissipating it only after a chase.

The amount of heat that cheetahs store during a sprint can be considerable, on the order of 60 times the heat produced at rest. Heat
storage raises the cheetahs body temperature. Cheetahs can therefore reach remarkable speeds but can sustain them only for short
durations and must rest for long periods of time between sprints. Because their bodies can support only so much heat storage,
cheetahs do not run when the temperature of their environments reaches approximately 105F. In other words, the amount of time a
cheetah runs and the distance it covers are limited by its body temperature.

Gazelles, common prey of cheetahs, have similar heat storage mechanisms and are similarly known for their speed and agility

How to dissipate heat?

Panting, sweating, and changes in blood flow.

3 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Metabolic Rate and Body Size: larger body sizes

have higher metabolic activity, and consume more
energy at rest, BUT the larger the organism, the
lower the metabolic rate/gram body wt..
In addition to activity level, an animals size influences its metabolic rate.

At rest, larger animals consume more energy and have higher metabolic rates
than smaller ones.

Resting, or basal, metabolic rate does not increase linearly with an animals
mass.

Measurements of resting metabolic rate in a wide range of organisms show

that metabolic rate increases with animal mass raised to the 3/4 power.

The larger the organism, the lower the metabolic rate per gram of body tissue.

This scaling pattern of cellular energy metabolism is remarkable because it holds across a diverse size range of unicellular and
multicellular organisms.

Metabolic Rate Affected by

Running & Body Size: larger
animals expend less
energy/unit body wt to cover a
distance
In the 1970s and 1980s, the American physiologist C. Richard
Taylor and his colleagues performed studies on the relationship
between metabolic rate and running speed in mammals of
different sizes.

Taylor and colleagues measured oxygen consumption during

running for a wide range of organisms.

Oxygen consumption was used as a measure of metabolic rate.

The researchers found that there is a linear increase in metabolic rate with speed in different-sized animals and that larger animals
expend less energy per unit body mass to move a given distance compared to smaller ones.

Similar studies were performed with kangaroos, which move by hopping rather than running.

Interestingly, it was found that it requires less energy to hop than to run.

This finding perhaps explains why kangaroos and other hopping animals survived, while many animals that run on four legs became
extinct in Australia 20,000 to 30,000 years ago with the arrival of early humans.

Studies have also compared the energetic costs of running on two legs and running on four legs, with the finding that the cost is the
same.

Hopping shown to be more energetically efficient than running.

Running on two legs energetically similar to running on four legs.

4 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Homeostasis and Thermoregulation: temperature affects chemical

reaction rates.
Metabolic rate is also affected by an animals internal body
temperature.

Temperature affects the rates of chemical reactions, which in turn

determine how fast fuel molecules can be mobilized and broken
down to supply energy for a cell.

Animals can be categorized by the sources of most of their heat.

Animals that produce most of their own heat as by-products

of metabolic reactions, including the breakdown of food, are
endotherms.

These animals usually, but not always,

maintain a constant body temperature
that is higher than that of their environment.

Mammals, including humans, and birds are endotherms.

Animals that obtain most of their heat from the environment are ectotherms. Ectotherms often regulate their body temperatures by
behavioral means such as moving into or out of the sun.

Most fish, amphibians, reptiles, and invertebrates are ectotherms.

Endotherms are sometimes referred to as warm-blooded and ectotherms as cold-blooded, but these terms are misleading
because, depending on environmental conditions, cold-blooded animals can have core body temperatures higher than warm-blooded
animals.

Endotherms have a higher metabolic rate than ectotherms.

As a result, they are able to be active over a broader range of external temperatures than ectotherms.

The activity level and metabolic rate of ectotherms both increase with increasing body temperature.

However, ectotherms have metabolic rates that are about 25% of endotherms of similar body mass and at similar body temperatures.

Although they can achieve activity levels similar to those of endotherms when their body temperatures are similar, ectotherms
cannot sustain prolonged activity.

The control of core body temperature, or thermoregulation, is a form of homeostasis and requires the coordinated activities of the
nervous, muscular, endocrine, circulatory, and digestive systems. A summary of how these systems work together in endotherms and
ectotherms is seen here.

Endotherms have higher energy consumption than ectotherms. Efficiency depends on the environment.

Salmon: dont eat during their trip upriver to spawn. Higher water temperatures cause them to metabolize
more rapidly than lower temps.

Can run out of stored energy too soon.

Energy Balance
Like core body temperature, blood-glucose levels, and blood pressure, the energy balance of an organism is often maintained at a
constant level.
5 Chapter 40 Animal Metabolism, Nutrition, and Digestion
An animal in energy balance takes in the same amount of
calories of energy from food that it uses over time to meet
its metabolic needs.

energy intake = sources of energy

energy use = ways in which energy is expended

For animals, the source of energy is the diet.

Energy use

Energy required for basic life processes (majority).

Depends on the level of physical activity.

Digestion and absorption of food from the diet itself requires energy.

When energy intake does not equal energy used, there is an energy imbalance. If an animal eats more food than it requires, energy
stores such as fat deposits grow over time.

During prolonged periods of an inadequate food supply or starvation, an animal consumes its own internal fuel reserves. Starvation
forces animals to deplete their glycogen and fat reserves first, and then, if no food is found, to resort to protein stores, primarily in
muscle tissue.

Excessive intake of food calories has led to an increasing and now critical public health problem: obesity.

Obesity is now an epidemic in many industrialized nations, including the United States, where about 36% of the adult
population is considered obese.

Obesity is a major public health concern because it increases the risk of diabetes, heart disease, and stroke and contributes
to a shorter life-span.

Why is obesity (more than 20% over ideal body wt) seen as a problem?

Increased risk of cardiovascular disease, type II diabetes, and of cancer. Decreased lifespan and quality of life.

Nearly 40% of North American adults are obese.

6 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Essential Amino Acids: (must come from diet) For Humans

(8 out of 20)
Animals metabolic pathways enable them to obtain energy from
the environment as well as to synthesize many of the compounds
needed to sustain life.

However, many nutrients necessary for life cannot be synthesized

by an animals metabolism and therefore must be acquired in the food that they eat.

An essential amino acid is one that cannot be synthesized by cellular biochemical pathways and instead must be ingested.

Most animals can synthesize about half of their amino acids.

Humans are unable to synthesize 8 of the 20 amino acids. We have to obtain these eight essential amino acids in our diets.

Minerals Required by Humans: chemical elements other than C, H, O and N that must
come from diet.
Dietary minerals are chemical elements other
than carbon, hydrogen, oxygen, and nitrogen
that are required in the diet and must be
obtained in the food that an animal eats.

They include such elements as calcium, iron,

phosphorus, potassium, zinc, and magnesium.

Calcium is required for building

skeleton, and iron in hemoglobin binds
oxygen and transports it in the blood.

Humans typically obtain the minerals

sodium and chloride, which ionize to
form salt crystals, through common
table salt and other foods.

Many wild animals seek exposed rock that they lick to obtain minerals and salts.
7 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Iron in diet

13 Essential Vitamins (organic

molecules needed in the diet) for
Humans
Vitamins are organic molecules that are required in very small amounts in the
diet.

Vitamins have diverse roles, some binding to and increasing the activity of
particular enzymes, others acting as antioxidants or chemical signals.

Thirteen essential vitamins have been identified for humans.

In humans as well as in other animals, vitamin deficiency can have serious

consequences.

Humans and other primates cannot synthesize vitamin C. Without ingesting sufficient vitamin C, humans develop scurvy, a
disease characterized by bleeding gums, loss of teeth, and slow wound healing.

Deficiencies of vitamins B1, B6, and B12 can cause nervous system disorders and various forms of anemia.

Vitamin D is essential for the absorption of calcium in the diet and thus to skeletal growth and health. With adequate
exposure to ultraviolet solar radiation, skin cells synthesize enough vitamin D to sustain a growing body. However, fairer-
skinned people inhabiting more northern regions of the world produce less vitamin D and therefore require more of it in their
diet.

A clear role for vitamin E remains uncertain, but its absence is often linked to anemia.

What are vitamins used for?

Assisting enzymatic function

Antioxidants.

Chemical signals.

What if you dont get enough vitamins in your diet?

Vitamin C deficiency: week connective tissue, scurvy, slow/poor wound-healing.

Vit B1, B6, and B12: nervous system disorders.

Vit D: impaired calcium absorption/skeletal issues. Yup UV stimulates your body to synthesize
vit D, but most of us do not get enough exposure to provide an adequate supply.

Vit E: anemia, reduced wound-healing.

8 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Filter Feeding: common in the animal kingdom,

aquatic environments. Convergent evolution.
Animals have evolved a diversity of ways to capture food and mechanisms for breaking down that
food before its digestion in the gut.

Suspension filter feeding

The most common form of food capture by animals

water with food suspended in it is passed through a sievelike structure.

possible only in aquatic environments, and it is ubiquitous there

evolved multiple times independently on many different branches of the tree of life, so
represents a form of convergent evolution.

Many worms and bivalve mollusks, such as scallops, clams, and oysters, pump water over their gills to
trap food particles suspended in the water.

Other aquatic organisms such as baleen whales move water with food particles suspended in it through filters in their oral cavity and then convey the
food into the gut.

Suction Feeding: rapid expansion of the mouth

cavity draws material in. Common in fish.
Many large aquatic animals capture their prey in other ways. Many fish feed by suction.

A rapid expansion of the fishs mouth cavity draws water and the desired prey into the
mouth.

After the fish closes its mouth, the water is pumped out of the mouth cavity past the
gills.

The prey is trapped inside the mouth, moves into the pharynx (part of the throat), and is
broken up by specialized pharyngeal jaws (a second set of jaws in the throat) before
being swallowed.

Suction feeding allows organisms to be sit-and- wait predators, hiding within a coral reef or under a rock before rapidly striking to capture prey moving
in front of them.

Many insects that bite to obtain a blood meal also rely on suction to draw the blood of their prey into their digestive system. Young mammals also feed
by suckling milk from their mothers breast, using their tongue and also generating suction as fish do.

Suction feeding is common in insects

Aphids

Assassin bug feeding on an aphid nymph

And of course, suction feeding is common in mammal infants even adults.

Active Swimming to Feed: using size and speed to

advantage. Need jaws.
Larger fish and marine mammals like sharks, whales, and dolphins actively swim to
capture their prey.

Their larger size and speed enables them to catch smaller or similar-sized prey.

The evolution of jaws and teeth considerably enhanced the ability of these and other vertebrate animals to capture
greater amounts of food, allowing them to lead a more active and energetically demanding lifestyle.
9 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Phylogeny of Vertebrates (Jaws) Jaws and teeth are significant adaptations, key to
vertebrate success.
Jaws and teeth were an important evolutionary
innovation for active predators.

Some of the first vertebrates, the jawed fishes,

became dominant in their aquatic environment
through the ability to swim and bite forcefully to
obtain their food.

Jawed fish evolved from jawless ancestors, and jaws

can be found in present-day fish, amphibians,
reptiles, and mammals.

Jaws are key to the evolutionary success of

vertebrates.

They are thought to have evolved from cartilage

that supported the gills, providing a spectacular
example of an organ adapted for one function (gill
support) changing over time to become adapted for
an entirely different function (predation).

Jaws and Teeth: mammals have the temporomandibular (enables the teeth
of upper and lower jaws to fit well) joint, and
huge variety of teeth adapted for different
functions.
Among vertebrates, mammals evolved a specialized jaw joint,
the temporomandibular joint, as well as a great diversity of
specialized forms of teeth.

The temporomandibular joint allowed the teeth of the lower and

upper jaws to fit together precisely.

This anatomy facilitated the specialization of teeth with cutting

and crushing surfaces, enabling mammals to break down a
variety of foods mechanically before swallowing.

Here, the arrangement of specialized teeth in mammals with different diets is illustrated.
10 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Organization of Digestive Tracts: mechanical and chemical

process. A long, one-way digestive tract enables specialization
and efficiency.
After animals obtain food, they break it down through the process of digestion.

Digestion requires that the food be isolated in a specialized compartment so that it can
be broken down chemically without damage to other cellular organelles or structures
of the body.

Many animals have digestive systems that allow the transport of food by a digestive
tube that runs from an animals mouth to its anus. It is known as the gut or digestive
tract.

Because the food is moved in a single direction, particular regions of the digestive
tract can be specialized for different functions: storage, chemical breakdown of
different kinds of food, absorption of released nutrients, and elimination of waste
products.

The digestive tracts of animals are commonly divided into a foregut, midgut, and
hindgut.

The foregut serves as an initial storage and digestive chamber.

The midgut is where the remainder of digestion and most nutrient absorption takes
place.

The hindgut is where water and minerals are reabsorbed, leaving the waste products, or feces

Swallowing: controlled by the autonomic nervous system after mechanical processing and
mixing with amylase.
Most animals jump-start digestion by breaking down
food mechanically.

Many animals break down food with the aid of jaws and
teeth.

Insects, which lack jaws and teeth, manipulate and

break down their food with mouthparts called
mandibles before it passes into their gut.

In mammals, food entering the mouth is mixed with

salivary secretions that contain amylase, an enzyme
that breaks down carbohydrates, to begin the digestion
of sugars and starches.

Mammals and other land vertebrates have a muscular tongue that facilitates food manipulation and transport within the mouth
cavity.

Swallowing, which is controlled by the autonomic nervous system, is a complex set of motor reflexes that involves several muscles
and structures in the rear of the mouth and the pharynx, the region of the throat that connects the nasal and mouth cavities. Once
initiated, swallowing reflexes occur without voluntary control.

Birds, alligators, crocodiles, and earthworms break down food into smaller pieces further along their digestive tracts in the gizzard, a
compartment with thick muscular walls.

Birds, alligators, crocodiles, and earthworms have a further mechanical digestion stage in the
gizzard.

Insects often use their mandibles to cut food into smaller chunks.
11 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Digestive Control: that oh so suggestible brain. Amino acids are absorbed

into the blood.
After being ingested and
mechanically broken down, the
food is transported into a
stomach or crop, one of the
main sites of protein
breakdown.

Hydrochloric acid (HCl) and

digestive enzymes that break
down proteins into amino acids,
which are then absorbed into
the bloodstream.

Cells secrete these digestive

enzymes in an inactive form;
otherwise, they would be
digested themselves. After
secretion, the inactive enzymes
are activated by a change in
their chemical structure.

The primary digestive enzyme

produced in the stomach is
pepsin, an enzyme that breaks
down proteins.

The cells lining the stomach also secrete a peptide hormone called gastrin in response to food in the stomach. Gastrin
stimulates the cells lining the stomach to increase their production of HCl. Gastrin secreted by the stomach is
absorbed within the small intestine and recirculated to the cells lining the stomach wall. There, the recirculated gastrin
stimulates the stomach lining cells to increase their production of gastrin, forming a positive feedback loop. The
feedback loop ensures that protein digestion occurs in response to the start of a meal as food enters the stomach. If
the pH of the stomach becomes too low, gastrin secretion is inhibited by negative feedback control.

The stomach walls contract to mix the contents of the stomach, aiding their digestion. Waves of muscular contraction,
called peristalsis, move the food toward the base of the stomach. There, the pyloric sphincter, a band of muscle, opens
and allows small amounts of digested food to enter the small intestine. Opening and closing of the pyloric sphincter
regulates the rate at which the stomach empties, allowing time for digestion and absorption of the food products
released into the small intestine.

Snakes and Large Whole Prey: months between meals. In humans, about 4 hours for the
stomach to empty.
Animals often eat large amounts of food in a short time, but it takes much longer
to digest that food.

In humans, it typically takes about four hours for the stomach to empty, allowing
digestion and absorption of nutrients to occur between meals. I

n other animals, stomach emptying can take much longer. Most carnivorous
animals consume large and infrequent meals, resulting in long periods of
digestion and nutrient absorption compared with humans.

Snakes such as pythons that engulf large prey whole elevate their metabolic rate
to high levels during digestion and spend several days digesting and absorbing the nutrients from their meal.

This requires an extensive remodeling of their gut to produce new secretory and absorptive cell surfaces to digest their
single large meal. These extra cells are not retained between meals because of the high energy cost of maintaining
them.
12 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Small Intestine: protein and

carbohydrate digestion
continues, and fat digestion
(Bile) starts.
After the food enters the small intestine,
protein and carbohydrate digestion
continue, and fat digestion begins. The
small intestine is made up of three
sections. The initial section is the
duodenum, into which the food enters
from the stomach. Most of the remaining
digestion takes place in the duodenum.
The next two sections, the jejunum and
ileum, carry out most nutrient
absorption.

Two accessory organs, the liver and

pancreas, aid in the digestion of proteins,
carbohydrates, and fats in the duodenum
of the small intestine. The liver produces
bile, which aids in fat digestion by
breaking large clusters of fats into
smaller lipid droplets, a process called
emulsification. Bile produced by the liver is stored in the gallbladder. When fats enter the duodenum, cells
lining the duodenum release a peptide hormone called cholecystokinin (CCK), which causes the gallbladder
to contract, thus releasing the bile into the duodenum.

The pancreas functions as both an endocrine gland, secreting hormones directly into the blood, and an
exocrine gland, secreting substances into ducts that connect to the duodenum. The pancreas produces a
variety of digestive enzymes, including lipase, which breaks down fats, and trypsin, which further breaks
down proteins. Like pepsin produced by the stomach, the pancreas produces trypsin in an inactive form
called trypsinogen to avoid digesting itself.

The pancreas also secretes bicarbonate ions, which neutralize the acid produced by the stomach. Enzymes
that break down proteins work best in acidic environments like that of the stomach, but most proteins, like
those that break down carbohydrates and fats, are denatured in acidic environments.
13 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Villi and Microvilli: increased surface area for absorption in the

jejunum and ileum.
Final digestion of proteins and carbohydrates occurs in the jejunum
and ileum. These two sections of the small intestine have highly
folded inner surfaces, called villi, and the cells constituting the villi
themselves have highly folded surfaces, called microvilli.

Together, villi and microvilli greatly increase the surface area for the
absorption of nutrients.

Cells that line the small intestine are connected by tight junctions.
These junctions force the products of digestion to be absorbed
across the microvilli surfaces, controlling their movement through
the cell and into the bloodstream, rather than leaking between the
cells.

The microvilli of the cells lining the small intestine secrete enzymes
that cleave peptides into amino acids, which can then be absorbed
across the plasma membrane. They also secrete enzymes that break sugars into their subunits, which can also be
readily absorbed.

Glucose Absorption:
transported across the
membrane (fats just
diffuse across the
membrane).
Nutrient molecules, such as
glucose and amino acids, are often
co-transported into cells lining the
intestine with sodium ions from
gut contents.

The sodium ion binds to a

transmembrane protein on the
surface of cells lining the intestine
that also binds the nutrient
molecule. Absorption of the
nutrient molecule into the cell is
driven by the movement of the
sodium ion down its concentration
gradient.

Because the products of fat

digestionfatty acids and glycerol
are lipid soluble, they do not
require a carrier protein for transport. Instead, they readily diffuse across the lipid plasma membrane, entering the
mucosal cell, where they are further broken down and transported to the bloodstream.
14 Chapter 40 Animal Metabolism, Nutrition, and Digestion

Large Intestine/colon: reabsorption of water and mineral ions. Too much

reabsorption = constipation; not enough = diarrhea.
After the absorption of nutrients in the small intestine, water
and mineral ions are reabsorbed in the large intestine, or
colon, of the hindgut.

By the time the gut contents reach the large intestine, the
nutrients have been absorbed into the body, but water and
inorganic ions remain.

These are absorbed in the large intestine until the contents

form semisolid feces.

Excess water absorption can cause constipation, whereas

too little water reabsorption results in diarrhea. Diarrhea and
constipation also can be caused by toxins released by
microorganisms, such as the bacterium Vibrio cholera, the
cause of cholera, in the food an animal eats.

Digestion is not a solo endeavor. Large populations of

bacteria reside in the small and large intestine and help extract nutrients that the animals body
cannot extract itself.

The principle gut resident is Escherichia coli.

The bacteria nourish themselves by aiding in the digestion of the hosts gut contents, but also provide
nutrients and certain vitamins, such as biotin and vitamin K, that the animal cannot produce itself.

Vitamin deficiency therefore can sometimes result from prolonged antibiotic medication that kills
large numbers of gut bacteria.

The mutual benefits to the host animal and the bacteria ensure the success of their symbiotic
relationship.

Mammalian Intestine: not a passive tube active, made up of specialized

layers of tissues.
The digestive tract is not a passive tube. It secretes enzymes and other
chemical compounds, absorbs nutrients, and actively moves food through
the body.

The digestive tract is made up of several layers of tissue, each with a

specialized function.

The central space, or lumen, through which the gut contents travel is
surrounded by an inner tissue layer, the mucosa, which has secretory and
absorptive functions. The cells of the mucosa secrete mucus to protect
the gut wall from digestive enzymes, and, in the stomach, hydrochloric
acid. Surrounding the mucosa is the submucosa, a layer containing blood
vessels, lymph vessels, and nerves.

Outside these layers are two smooth muscle layers.

An inner circular muscle layer contracts to reduce the size of the

lumen.

An outer longitudinal muscle layer contracts to shorten small sections of the gut.

These two muscle layers contract alternately to mix gut contents and to move the contents along the digestive tract from
compartment to compartment.
15 Chapter 40 Animal Metabolism, Nutrition, and Digestion
Between the two muscle layers are autonomic nerves that control the contractions of the two sets of smooth muscle.

An outer layer of cells and connective tissue called the serosa covers and protects the gut.

The gut is supported in the abdominal cavity by a membrane called the mesentery, through which blood vessels, nerves,
and lymph travel to supply the gut.

ForeGut Fermenter in cows, sheep: improved

digestion of cellulose
Herbivorous animals evolved specialized digestive tracts that enhance their ability to digest
cellulose and other plant compounds.

Most herbivores, such as cows and termites, lack cellulase, the enzyme that breaks down cellulose.

Instead, they have specialized compartments in their digestive tract that contain large bacterial
populations, which do produce cellulase.

This association of herbivores and their gut microbes is another example of a symbiotic relationship.

Rather than the single stomach of other mammals, ruminants (cattle, sheep, and goats) have a four-
chambered stomach that is highly specialized to enhance the ability of their gut bacteria to digest
plants.

The first two chambersthe rumen and the reticulum harbor large populations of anaerobic
bacteria that break down cellulose by fermentation, and some of the
products are used as nutrients by the host. Carbon dioxide and methane gas
are also produced as a result of bacterial fermentation.

Actually, it is bacteria in the g.i. which produce the cellulose

which digests the cellulose.

HindGut Fermenters: less efficient hence the rabbit habit of eating its
feces.
In contrast to the foregut fermentation of cows and sheep, other mammalian herbivores, such as
koalas, rabbits, and horses, digest the plant material they eat by hindgut fermentation.

Hindgut fermentation occurs in the colon and in the cecum, a chamber that branches off the
large intestine. Because the fermentation products released from the cecum of a koala or horse
have already passed through the small intestine, the main site of nutrient absorption, hindgut
fermentation is less efficient than foregut fermentation in terms of nutrient extraction.