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In humans, earwax lubricates the outer ear canal and protects the
eardrum. Honeybees use a wax secreted by glands in their abdo-
men to construct the comb in which larvae are raised and honey
is stored (Figure .A).
Many plants secrete waxes that form a protective exterior
layer, which greatly reduces water loss from the plants and resists
invasion by infective agents such as bacteria and viruses. is
waxy covering gives cherries, apples, and many other fruits their
shiny appearance (Figure .B).
Phospholipids Provide the Framework
of Biological Membranes
Phosphate-containing lipids called phospholipids are the pri-
mary lipids of cell membranes. In the most common phospho-
lipids, glycerol forms the backbone of the molecule as in tri-
glycerides, but only two of its binding sites are linked to fatty
acids (Figure .). e third site is linked to a polar phosphate
group, which binds to yet another polar unit. e end of the
Fats, Cholesterol, and Coronary
Butter! Bacon and eggs! Ice cream! Cheese-
cake! Possibly you think of such foods as irre-
sistible, off limits, or both. After all, who
doesn’t know about animal fats, cholesterol,
and hardening of the arteries? Hardening of
the arteries, or atherosclerosis, is a condition
in which deposits of lipid and ﬁbrous material
called plaque build up in the walls of arteries,
the vessels that supply oxygenated blood to
body tissues. Plaque reduces the internal di-
ameter of the arteries, restricting or even
completely blocking the ﬂow of blood. Block-
age of the coronary arteries that supply oxy-
genated blood to the heart muscle (see ﬁgure)
can severely impair heart function, a condition
called coronary heart disease. In extreme
cases, it can lead to destruction of heart mus-
cle tissue, as occurs in a heart attack (myocar-
Your body requires a certain amount of
cholesterol, but the liver normally makes
enough to meet this demand. Additional
cholesterol is made from fats taken in as
food. Cholesterol is found in the blood
bound to low-density lipoprotein (LDL) and
high-density lipoprotein (HDL). LDL choles-
terol is considered “bad” because clinical
studies have shown a positive correlation be-
tween its level in the blood and the risk for
coronary heart disease. LDL cholesterol con-
tributes to plaque formation as atherosclero-
sis proceeds. In contrast, HDL cholesterol is
“good” because clinical studies have shown
that high levels of this form appear to pro-
vide some protection against coronary heart
disease. Simplifying, HDL cholesterol re-
moves excess cholesterol from plaques in ar-
teries, thereby reducing plaque buildup. The
cholesterol that has been removed is trans-
ported by the HDL cholesterol to the liver
where it is broken down.
Fats in food affect cholesterol levels in the
blood. Diets high in saturated fats raise LDL
cholesterol levels, but levels of HDL choles-
terol appear not to be affected by such a diet.
Foods of animal origin typically contain satu-
rated fats, and foods of plant origin typically
contain unsaturated fats.
In the food industry, unsatu-
rated vegetable oils are often pro-
cessed to solidify the fats. The pro-
cess, partial hydrogenation, adds
hydrogen atoms to unsaturated
sites, eliminating many double
bonds and generating substances
known as trans fatty acids (or trans
fats). Usually the hydrogen atoms
at a double bond are positioned on
the same side of the carbon chain,
producing a cis (Latin, “on the
same side”) fatty acid:
but in a trans (Latin, “across”) fatty
acid, the hydrogen atoms are on
different sides of the chain at some double
Trans fatty acids are found in many vegetable
shortenings, some margarines, cookies,
cakes, doughnuts, and other foods made with
or fried in partially hydrogenated fats.
Research from human feeding studies has
shown that trans fatty acids raise LDL choles-
terol levels nearly as much as saturated fatty
acids do. More seriously, intake of trans fatty
acids at levels found in a typical U.S. diet also
appears to reduce HDL cholesterol levels. In
addition, clinical studies have demonstrated a
positive correlation between the intake of
trans fatty acids and the occurrence of coro-
nary heart disease. A regulation to add the
trans fatty acid content to nutritional labels
went into effect in the United States in January
. A number of federal and state agencies
and cities have or are considering legislation
to ban trans fatty acids in food. For instance,
in , California became the ﬁrst state to
ban trans fatty acids in restaurants, following
a number of cities around the country.
One of the many unanswered questions
about dietary cholesterol is why some popula-
tions can consume large quantities of fatty
foods of the “wrong” kind yet rarely develop
atherosclerosis. For example, atherosclerosis
was once virtually nonexistent in Inuits,
whose diet in their native culture contained
more than % animal fat; however, athero-
sclerosis developed in that same population
when they adopted a “civilized” diet and life-
style. In France, the incidence of atherosclero-
sis is relatively low even though cheese and
other dairy products are diet staples. Of
course, the French say that wine keeps them
FOCUS ON Applied Research
Atherosclerotic plaques (bright areas) in the coronary arteries
of a patient with heart disease.
UNI T SI X ANI MAL STRUCTURE AND FUNCTI ON 966
When you sit without moving for long periods of time, as
you might during an airline ight, the lack of skeletal muscular
activity greatly reduces the return of venous blood to the heart.
As a result, the blood pools in the veins of the body below the
heart, making the hands, legs, and feet swell. e motionless
blood can also form clots, particularly in the veins of the legs, a
condition called deep vein thrombosis. Deep vein thrombosis
oen does not cause symptoms, but can cause serious medical
problems if a clot breaks loose and moves elsewhere in the body,
such as to the lungs. Raising the arms and getting up at intervals
to exercise or contracting and relaxing the leg muscles as you sit
can relieve this condition.
Disorders of the Circulatory System Are
Major Sources of Human Disease
e layer of endothelial cells lining the arteries and veins is nor-
mally smooth and does not impede blood ow. However, several
conditions, including bacterial and viral infections, chronic hy-
pertension, smoking, and a diet high in fats, can damage the en-
dothelial cells, exposing the underlying smooth muscle tissue,
which begins a cycle of injury and repair leading to lesions.
(Focus on Applied Research in Chapter 3 discussed the relation-
ship between fats and cholesterol and coronary artery disease,
and the Unanswered Question in Chapter 3 discussed how the en-
zymes that synthesize fatty acids and cholesterol are regulated in
cells.) ickened deposits of material called atherosclerotic plaques
may form at the damaged sites (Figure 42.1I). e plaques, which
consist of cholesterol-rich fatty substances, smooth muscle cells,
and collagen deposits, reduce the diameter of the blood vessel and
impede blood ow. Worse, the damaged endothelial lining may
stimulate platelets to adhere and trigger the formation of blood
clots. e clots further reduce the vessel diameter and ow and
may break loose, along with segments of plaque material, to block
ner vessels in other regions of the body.
Atherosclerosis has its most serious eects in the smaller ar-
teries of the body, particularly in the ne coronary arteries that
serve the heart muscle. Here, the plaques and clots reduce or
block the ow of blood to the heart muscle cells. Serious blockage
can cause a heart attack—the death of cardiac muscle cells de-
prived of blood ow. e blockage of arteries in the brain by
plaque material or blood clots released from atherosclerotic ar-
teries is also a common cause of stroke—a loss of critical brain
functions because of the death of nerve cells in the brain. Heart
attacks and strokes are the most common causes of death in
North America and Europe.
e risk of heart attacks and stroke can be reduced by avoid-
ing the conditions that damage the blood vessel endothelium. A
diet low in fats and cholesterol, avoidance of cigarette smoke,
and a program of exercise can reduce epithelial damage and
plaque deposition. Medication and exercise, or exercise alone,
can also reduce the eects of hypertension. ere are good indi-
cations that these preventive programs can also reduce the size
of existing atherosclerotic plaques.
STUDY BREAK 42.4 <
1. How is blood ﬂow through capillary networks controlled?
2. Explain how, in contrast to most body tissues, the brain does not
allow exchange of molecules and ions with blood.
3. Describe the two major mechanisms that drive the exchange of
molecules and ions between the capillaries and the interstitial ﬂuid.
42.5 Maintaining Blood Flow
Arterial blood pressure is the principal force moving blood to the
tissues. Blood pressure must be regulated carefully so that the
brain and other tissues receive adequate blood ow, but not so
high that the heart is overburdened, risking damage to blood ves-
sels. e three main mechanisms for regulating blood pressure
are controlling cardiac output (the pressure and amount of blood
pumped by the le and right ventricles), the degree of constric-
tion of the blood vessels (primarily the arterioles), and the total
blood volume. e sympathetic division of the autonomic ner-
vous system and the endocrine system interact to coordinate
Cardiac Output Is Controlled by Regulating the
Rate and Strength of the Heartbeat
Regulation of the strength and rate of the heartbeat starts at
stretch receptors called baroreceptors (a type of mechanorecep-
tor; see Section 39.2), located in the walls of blood vessels. By
detecting the amount of stretch of the vessel walls, barorecep-
tors constantly provide information about blood pressure. e
baroreceptors in the cardiac muscle, the aorta, and the carotid
arteries (which supply blood to the brain) are the most crucial.
Signals sent by the baroreceptors go to the medulla within the
brain stem. In response, the brain stem sends signals via the au-
tonomic nervous system that adjust the rate and force of the
heartbeat: the heart beats more slowly and contracts less force-
fully when arterial pressure is above normal levels, and it beats
faster and contracts more forcefully when arterial pressure is
below normal levels.
A. Normal artery B. Clogged artery
Wall of artery,
Atherosclerosis. (A) A normal coronary artery. (B) A coronary artery that is
partially clogged by an atherosclerotic plaque.
CHAPTER ANIMAL NUTRITION 1021
moisten the mixture before it enters the esophagus and passes into
the crop. ese secretions begin the process of chemical digestion.
From the crop, the food mass enters the gizzard, which grinds it
into smaller pieces. ese food particles enter the stomach, in
which food is stored and digestion begins. Insect stomachs have
saclike outgrowths, the gastric ceca (caecus ϭ blind), where en-
zymes hydrolyze the digestive contents; the products of digestion
are absorbed through the walls of the ceca. e undigested contents
then move into the intestine for further digestion and absorption.
At the end of the intestine, water is absorbed from the undigested
matter and the remnants are expelled through the anus. e diges-
tive systems of other arthropods are similar to the insect system.
DIGESTION IN A BIRD A pigeon (Figure 45.4C ) picks up seeds
with its bill. e bird’s tongue moves the seeds into its mouth,
where they are moistened by mucus-lled saliva and swallowed
whole (birds have no teeth). e seeds then pass through the
pharynx into the esophagus. (In some cases, birds crack open
seeds with their bills and ingest seed kernels in a similar fashion.)
e anterior end of the esophagus is tubelike; at the posterior end
is the pouchlike crop, in which the bird can store large quantities
of food. From the crop, the food passes into the anterior glandular
portion of the stomach, called the proventriculus, which secretes
digestive enzymes and acids. e posterior end is the gizzard, in
which the seeds are ground into ne particles, aided by ingested
bits of sand and rock. e food particles are released into the in-
testine, where the liver secretes bile and the pancreas adds diges-
tive enzymes. e molecular subunits produced by enzymatic di-
gestion are absorbed as the mixture passes along the intestine,
and the undigested residues are expelled through the anus.
Many of the structures of the pigeon’s digestive system, in-
cluding the mouth, pharynx, esophagus, stomach, intestine,
liver, and pancreas, occur in almost all vertebrates.
STUDY BREAK 45.2 <
1. Distinguish between intracellular digestion and extracellular
2. What are the ﬁve steps of food processing in a digestive tube?
THINK OUTSIDE THE BOOK
Use the Internet or a research paper search tool such as
PubMed (http://www.ncbi.nlm.nih.gov/PubMed/) to list the
research question being addressed in three projects involving
bird digestive systems.
45.3 Digestion in Humans
and Other Mammals
Mammals digest foods using the same ve steps as other animals
with a digestive tube: mechanical processing, secretion of en-
zymes and other digestive aids, enzymatic hydrolysis, absorption
of molecular subunits, and elimination. e mammalian diges-
tive system is a series of specialized digestive regions that perform
these steps, including the mouth, pharynx, esophagus, stomach,
small and large intestines, rectum, and anus (Figure 45.5). ese
regions are under the control of the nervous and endocrine sys-
tems. e digestive system is not identical in all mammals. Spe-
cic adaptations have produced variants of the digestive system
such as occurs in humans. An example is the digestive system of
ruminants (discussed in Section 45.5).
Humans Require Speciﬁc Essential Amino
Acids, Fatty Acids, Vitamins, and Minerals
in Their Diet
e human digestive system meets our basic needs for fuel mole-
cules and for a wide range of nutrients, including the molecular
building blocks of carbohydrates, lipids, proteins, and nucleic
acids. If the diet is adequate, the digestive system also absorbs the
essential nutrients—the amino acids, fatty acids, vitamins, and
minerals that cannot be synthesized within our bodies.
ESSENTIAL AMINO ACIDS AND FATTY ACIDS ere are eight
essential amino acids for adult humans: lysine, tryptophan, phe-
nylalanine, threonine, valine, methionine, leucine, and isoleu-
cine. Infants and young children also require histidine. e pro-
teins in sh, meat, egg whites, milk, and cheese supply all the
essential amino acids, provided those foods are eaten in adequate
quantities. In contrast, the proteins of many plants are decient in
one or more of the essential amino acids. Corn, for example, con-
tains inadequate amounts of lysine, and beans contain little me-
thionine. Vegetarians, and especially vegans who eat a diet with
no animal-derived nutrients, must choose their foods carefully to
obtain all of the essential amino acids (Figure 45.6). Such diets
typically include combinations of foods, each of which provides
some amino acids, and that together contain all of the essential
amino acids. An example is including in the diet rice or corn or
other grains (low in lysine but high in methionine) with lentils or
soybeans (perhaps in the form of tofu) or other legumes (low in
methionine but high in lysine).
If the diet lacks one or more essential amino acids, many en-
zymes and other proteins cannot be synthesized in su cient
quantities. e resulting protein deciency is most damaging to
the young, who must rapidly synthesize proteins for develop-
ment and growth. Even mild protein starvation during preg-
nancy or for some months aer birth can retard a child’s mental
and physical development.
Only two fatty acids, linoleic acid and linolenic acid, are es-
sential in the human diet. Both are required for synthesis of
phospholipids forming parts of biological membranes and cer-
tain hormones. Because almost all foods contain these fatty
acids, most people have no problem obtaining them. However,
people on a low-fat diet that is decient in linoleic acid and lino-
lenic acid are at serious risk for developing coronary heart dis-
ease. at is, there is an inverse correlation between the concen-
tration of these essential fatty acids in the diet and the incidence
of coronary heart disease. is is illustrated in the case of Hindu
UNI T SI X ANI MAL STRUCTURE AND FUNCTI ON
VITAMINS Humans require 13 known vitamins in their diet.
Many metabolic reactions depend on vitamins, and the absence
of one vitamin can aect the functions of the others. ese essen-
tial nutrients fall into two classes: water-soluble (hydrophilic)
vitamins and fat-soluble (hydrophobic) vitamins (summarized
in Table .). e body stores excess fat-soluble vitamins in adi-
pose tissues, but any amount of water-soluble vitamins above
daily nutritional requirements is excreted in the urine. us,
meeting the daily minimum requirements of water-soluble vita-
mins is critical for good health. e body can tap its stores of
fat-soluble vitamins to meet daily requirements, however, these
stores are quickly depleted so prolonged deciencies of the fat-
soluble vitamins also negatively aect health.
Most of you get all the vitamins you need through a normal
and varied diet that includes meats, sh, eggs, cheese, and vege-
tables. Vitamin supplements are usually necessary only for strict
Muscular sac; stretches to store food;
secretes mucus and gastric juice that
contains pepsinogen, the precursor to
the protein-digesting enzyme pepsin,
and hydrochloric acid (HCI).
Duodenum receives secretions from
liver, gallbladder, and pancreas.
Produces enzymes that complete
digestion of proteins, carbohydrates,
and nucleic acids; absorbs products
Absorbs water and mineral ions;
secretes mucus and bicarbonate
ions; concentrates undigested
matter into feces.
Stores feces; distension stimulates
expulsion of feces.
End of system; opening through
which feces are expelled.
Secretes enzymes (proteases, amylases,
lipases, nucleases) that break down all
major food molecules and bicarbonate
ions that neutralize digestive contents.
Muscular contractions move food
to esophagus by swallowing reﬂex.
Mouth (oral cavity)
Entrance to system; food is
moistened and chewed;
polysaccharide digestion starts.
Stores and concentrates bile
secreted by liver.
Secretes bile, which emulsiﬁes
fats, and bicarbonate ions.
Muscular, mucus-moistened tube
moves food from pharynx
Secrete saliva, which contains
lubricating mucus, amylase
(a starch-digesting enzyme),
lysozyme (an enzyme that kills
bacteria), and bicarbonate ions.
The human digestive system.
Rice, corn, or
(for example, tofu),
or other legumes
Eight essential amino acids
amino acids in a
vegetarians from India. eir diet consists mainly of low-fat
grains and legumes—clearly a low-fat diet—yet their rate of cor-
onary heart disease is higher than that in the United States and
Europe, where dietary fat content is higher.
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