Campbell Biology 12e (1) - 921-946

Unit 7 ANIMAL FORM AND
FUNCTION
Meet the “phage wrangler,” Steffanie Mexican border. In 2017, extreme personal circum-
Strathdee, Professor of Medicine in the Division stances prompted a radical shift in her research, as
of Infectious Diseases and Global Public Health detailed in The Perfect Predator: A Scientist’s Race
at the University of California, San Diego. Born in to Save Her Husband from a Deadly Superbug, a
Canada, she completed her M.Sc. and Ph.D. degrees book co-authored by Dr. Strathdee and her hus-
in epidemiology at the University of Toronto. Since band. In 2018, Dr. Strathdee became co-director of
joining the faculty at UC San Diego in 2004, she the newly established Center for Innovative Phage
has focused on HIV research and prevention in the Applications and Therapeutics at UC San Diego.
underserved populations of Tijuana, just across the
AN INTERVIEW WITH of which epidemiology is one part. the bacteria in the process. I emailed Tom’s
Epidemiology involves studying risk fac- doctor, who is a colleague of Tom’s and
Steffanie Strathdee tors and patterns, not just at an individual mine, and he thought the idea was worth a
level, like what people eat and how they shot. So, I had to go out and find people who
Tell us about your start in science. behave, but also the social, political, had phages that were active against Tom’s
I always had a natural curiosity as a young and economic forces that drive those bacterium, and that felt harder than looking
girl. I did not have an innate ability at science behaviors. for a needle in a haystack. Luckily, labs at the
or math, even though I was interested in Naval Medical Research Center in Maryland
them. I struggled in math. Even in university, What changed your research focus? and at Texas A&M University dedicated
I got D’s in calculus. I still don’t like math, but My husband became ill with a superbug themselves to the search and found phages
I’ve learned to surround myself with people infection—an infection caused by a bacte- that matched. The phages were grown, puri-
that do really well in math. I think it’s impor- rium that is resistant to many antibiotics. fied, and injected into Tom’s body, a billion
tant for students to realize that just because As an infectious disease epidemiologist, phages per dose. Three days later, Tom woke
they’re not good at everything doesn’t you would think that I would have un- up from his coma.
mean that they can’t pursue their dreams. derstood the global crisis we are facing in
multidrug-resistant organisms. But it wasn’t Tell us a bit more about your
How did you become interested in until it hit me on a personal level that I fully collaborators.
epidemiology? understood the threat. A Ph.D. student at Texas A&M worked
After both my course instructor and my around the clock and found the phages
How did the story of your husband’s that made up the first infusion. This
Masters and Ph.D. advisor passed away
from AIDS, I decided I wanted to focus on illness unfold? student was at a really low point in her
ending the HIV epidemic. I had started My husband and I were in Egypt, and he studies, thinking, “I don’t know what I’m
out in the laboratory and realized I was became very ill. The doctor gave him IV going to do with my career, I don’t know
really lousy at tissue culture experiments. antibiotics and said, “He’ll be fine,” but he if what I do really matters.” And then she
So I turned my attention to public health, wasn’t. It turned out that a gallstone had ended up helping save the life of a total
blocked his bile duct, stranger, and it’s jump-
causing an abscess started a whole new
. Dr. Steffanie Strathdee’s husband,
(cavity) to form, and
“This student . . . ended field—phage therapy.
Dr. Tom Patterson, holding an electron
micrograph of his superbug (left), and
a superbug moved up helping save the
into that abscess and What advice do you
Dr. Strathdee, with an artist’s rendition of
multiplied. Tom’s par- life of a total stranger, have for students
the bacteriophage that defeated it (right).
ticular bacterium had
51 different antibiotic-
and it’s jump-started starting to study
resistance genes. It was a whole new field.” biology?
winning its battle against Students should look for
the immune system, none mentors with whom they
of the standard antibiotic treatments were share the same values. For me, that was
working, and my husband was dying. I did what really turned my D in calculus and
a literature search and found a hundred- some of the troubles that I had in science
year-old, forgotten cure based on bacte- into success. I found mentors who were
riophages. These viruses, called phages for encouraging and helped me identify my
short, attack bacteria, but not human cells. strengths. Knowing what your weak-
Each phage is specific for a particular bacte- nesses and your strengths are is, I think,
rium. Phages inject their DNA into bacteria, one of the big components to making a
turning them into phage factories and killing successful career.
40
KEY CONCEPTS
Basic Principles of Animal
Form and Function
40.1 Animal form and function

are correlated at all levels of
organization p. 874
40.2 Feedback control maintains the

internal environment in many
animals p. 881
40.3 Homeostatic processes for

thermoregulation involve form,
function, and behavior p. 884
40.4 Energy requirements are related

to animal size, activity, and
environment p. 889
Study Tip Figure 40.1 Emperor penguins (Aptenodytes forsteri) live in Antarctica, Earth’s
coldest and windiest continent. In summer, these birds catch fish by diving down
Draw a diagram: When you encounter 500 meters in water only 2°C above freezing. In winter, the females forage and the
an example in the chapter of how an males incubate eggs while temperatures drop to –40°C and winds gust to 200 km/hr.
animal maintains a steady internal
state, draw a simple circuit diagram
(see example—illustrations are How do animals regulate their internal state even in
optional!). Label the variable being
controlled, a perturbation that affects
changing or harsh environments?
the variable, the response, and its effect Adaptations in form, function, and behavior help maintain an animal’s internal
in restoring the normal state. environment. Adaptations that limit variation in temperature and other internal
variables are widespread and diverse. Consider, for example, three adaptations
Variable: BODY TEMPERATURE that help an Emperor penguin stay warm:
(example: penguins)
Normal body Form (anatomy): An insulating
temperature
layer of fat (blubber) reduces heat loss
Perturbation: from most of the penguin’s body
Effect: body cold weather (blue body areas in this thermal Function (physiology): Rapid
temperature causes drop image). cycles of muscle contraction and
warms toward in body relaxation during
normal temperature shivering produce
heat at a cellular
level.
Response:
penguins huddle
together, reducing
exposed surface area
Go to Mastering Biology
For Students (in eText and Study Area)
• Get Ready for Chapter 40
• Figure 40.17 Walkthrough: Behavior: By packing together in
Thermoregulation in Humans groups of up to several thousand,
Emperor penguins greatly reduce
For Instructors to Assign (in Item Library) their exposure to wind and cold.
• Everyday Biology: How to Keep
Your Cool
• Tutorial: Thermoregulation
873
CONCEPT 40.1 example of convergent evolution (see Concept 22.3).
Natural selection often results in similar adaptations when
Animal form and function diverse organisms face the same environmental challenge,
such as overcoming drag during swimming.
are correlated at all levels Physical laws also influence animal body plans with regard
of organization to maximum size. As body dimensions increase, thicker skel-
etons are required to maintain adequate support. This limita-
Over the course of its life, an Emperor penguin faces the same
tion affects internal skeletons, such as those of vertebrates, as
fundamental challenges as any other animal, whether hydra,
well as external skeletons, such as those of insects and other
hawk, or human. All animals must obtain nutrients and oxygen,
arthropods. In addition, as bodies increase in size, the muscles
fight off infection, and survive to produce offspring. Given that
required for locomotion must represent an ever-larger fraction
all animal species share these and other basic requirements, why
of the total body mass. At some point, mobility becomes lim-
does their form, including anatomy—biological structure—
ited. By considering the fraction of body mass in leg muscles
vary so widely? The answer lies in natural selection and adapta-
and the effective force such muscles generate, scientists can
tion. Natural selection favors those variations in a population
estimate maximum speed for a wide range of body plans. In the
that increase relative fitness (see Concept 23.4). The evolution-
case of the 6-meter-tall dinosaur Tyrannosaurus rex, there is con-
ary adaptations that enable survival vary among environments
troversy, with some scientists calculating a top running speed as
and species but frequently result in a close match of form to
fast as that of an Olympic sprinter—30 km/hr (19 miles/hour),
function, as illustrated for the Emperor penguins in Figure 40.1.
but others inferring that T. rex was at best a fast walker.
Because structure and function are correlated, examining anat-
omy often provides clues to physiology—biological function.
An animal’s size and shape are fundamental aspects of form
Exchange with the Environment
that significantly affect the way the animal interacts with its
Animals must exchange nutrients, waste products, and gases
environment. Although we may refer to size and shape as
with their environment, and this requirement imposes an addi-
elements of a “body plan” or “design,” this does not imply a
tional limitation on body plans. Exchange occurs as substances
process of conscious invention. The body plan of an animal
dissolved in an aqueous solution move across the plasma mem-
is the result of a pattern of development programmed by the
brane of each cell. A single-celled organism, such as the amoeba
genome, itself the product of millions of years of evolution.
in Figure 40.3a, has a sufficient membrane surface area in con-
tact with its environment to carry out all necessary exchange.
Evolution of Animal Size and Shape In contrast, an animal is composed of many cells, each with
EVOLUTION Many different body plans have arisen during its own plasma membrane across which exchange must occur.
the course of evolution, but these variations fall within certain The rate of exchange is proportional to the membrane surface
bounds. Physical laws that govern strength, diffusion, move- area involved in exchange, whereas the amount of material that
ment, and heat exchange limit the range of animal forms. must be exchanged is proportional to the total body volume.
As an example of how physical laws A multicellular organization therefore works only if
constrain evolution, let’s consider every cell has access to a suitable aqueous environ-
. Figure 40.2 Convergent
how some properties of water limit the evolution in fast
ment, either inside or outside the animal’s body.
possible shapes for animals that are swimmers. Many animals with a simple internal organiza-
fast swimmers. Water is about 1,000 tion have body plans that enable direct exchange
times denser than air and also far more between almost all their cells and the external
viscous. Therefore, any bump on an environment. For example, a pond-dwelling
animal’s body surface that causes drag hydra has a saclike body plan and a body wall only
Seal
impedes a swimmer more than it would two cell layers thick (Figure 40.3b). Because its gas-
a runner or flyer. Tuna and other fast trovascular cavity opens to the external environ-
ray-finned fishes can swim at speeds ment, both the outer and inner layers of cells are
up to 80 km/hr (50 miles/hour). Sharks, constantly bathed by pond water. Another com-
penguins, dolphins, and seals are also mon body plan that maximizes exposure to the
relatively fast swimmers. As illustrated Penguin surrounding medium is a flat shape. Consider, for
by the three examples in Figure 40.2, instance, a parasitic tapeworm, which can reach
these animals all have a shape that is several meters in length (see Figure 33.11). A thin,
fusiform, meaning tapered on both flat shape places most cells of the worm in direct
ends. The similar streamlined shape contact with its particular environment—the
found in these speedy vertebrates is an Tuna nutrient-rich intestinal fluid of a vertebrate host.
874 UNIT SEVEN Animal Form and Function

. Figure 40.3 Direct exchange with the environment. hundreds of thousands of times smaller than that for a water
flea. Nevertheless, every cell in the whale has access to oxygen,
nutrients, and other resources. How is this accomplished?
Mouth
In whales and most other animals, the evolutionary adap-
Gastrovascular tations that enable sufficient exchange with the environ-
cavity
ment are specialized surfaces that are extensively branched
Exchange or folded (Figure 40.4). In almost all cases, these exchange
Exchange
surfaces lie within the body, an arrangement that protects
their delicate tissues from abrasion or dehydration and allows
for streamlined body contours. The branching or folding
Exchange greatly increases surface area (see Figure 33.8). In humans, for
example, the exchange surfaces for digestion, respiration, and
circulation each have an area more than 25 times larger than
that of the skin.
0.1 mm 1 mm
Internal body fluids link exchange surfaces to body cells.
(a) An amoeba, a single-celled (b) A hydra, an animal with The spaces between cells are filled with fluid, known in many
organism two layers of cells animals as interstitial fluid (from the Latin for “stand
between”). Complex body plans also include a circulatory
fluid, such as blood. Exchange between the interstitial fluid
Our bodies and those of most other animals are composed and the circulatory fluid enables cells throughout the body to
of compact masses of cells, with an internal organization obtain nutrients and get rid of wastes (see Figure 40.4).
much more complex than that of a hydra or a tapeworm. For Complex body plans offer numerous benefits. For exam-
such a body plan, increasing the number of cells decreases the ple, an external skeleton can protect against predators, and
ratio of outer surface area to total volume. As an extreme com- sensory organs can provide detailed information on the ani-
parison, the ratio of outer surface area to volume for a whale is mal’s surroundings. Internal digestive organs can break down
c Figure 40.4 Internal exchange surfaces of EXTERNAL ENVIRONMENT

complex animals. Most animals have surfaces that CO2
are specialized for exchanging chemicals with the Food O2
surroundings. These exchange surfaces are Mouth
usually internal but are connected to the ANIMAL
environment via openings on the body surface BODY
(the mouth, for example). The exchange surfaces
250 om
are finely branched or folded, giving them a Respiratory
very large area. The digestive, respiratory, and od
system
Blo
excretory systems all have such exchange

A microscopic view of the lung
surfaces. Chemicals exchanged across reveals that it is much more
these surfaces are transported throughout sponge-like than balloon-like. This
the body via the circulatory system. Heart construction provides an expansive
Cells
VISUAL SKILLS Using this diagram, explain wet surface for gas exchange with
how exchange carried out by animals can be
the environment (SEM).
described as both internal and external.
Nutrients Circulatory
system
Interstitial
fluid
Digestive Excretory
system system
100 om
50 om
The lining of the small intestine Anus Within the kidney, blood is filtered
has finger-like projections that across the surface of long, narrow
expand the surface area for Unabsorbed Metabolic waste products blood vessels packed into
nutrient absorption (SEM). matter (feces) (nitrogenous waste) ball-shaped structures (SEM).
CHAPTER 40 Basic Principles of Animal Form and Function 875

food gradually, controlling the release of stored energy. In but also regulates the level of sugar in the blood as a vital part
addition, specialized filtration systems can adjust the compo- of the endocrine system.
sition of the internal fluid that bathes the animal’s body cells. Just as viewing the body’s organization from the “bottom
In this way, an animal can maintain a relatively stable inter- up” (from cells to organ systems) reveals emergent properties, a
nal environment despite the fact that it is living in a change- “top-down” view of the hierarchy reveals the multilayered basis
able external environment. A complex body plan is especially of specialization. Organ systems include specialized organs
advantageous for animals living on land, where the external made up of specialized tissues and cells. Consider the human
environment may be highly variable. digestive system: Each organ has specific roles. In the case of
the stomach, one role is to initiate protein breakdown. This
process requires a churning motion powered by stomach mus-
Hierarchical Organization of Body Plans cles, as well as digestive juices secreted by the stomach lining.
Cells form a working animal body through their emergent Producing digestive juices, in turn, requires highly specialized
properties, which arise from successive levels of structural and cell types: One cell type secretes a protein-digesting enzyme, a
functional organization (see Concept 1.1). Cells are organized second generates concentrated hydrochloric acid, and a third
into tissues, groups of cells with a similar appearance and produces mucus, which protects the stomach lining.
a common function. Different types of tissues are further The specialized and complex organ systems of animals are
organized into functional units called organs. (The simplest built from a limited set of cell and tissue types. For example,
animals, such as sponges, lack organs or even true tissues.) lungs and blood vessels have different functions but are lined
Groups of organs that work together, providing an additional by tissues that are of the same basic type and therefore share
level of organization and coordination, make up an organ many properties.
system (Table 40.1). Thus, for example, the skin is an organ There are four main types of animal tissues: epithelial,
of the integumentary system, which protects against infec- connective, muscle, and nervous. Figure 40.5 explores the
tion and helps regulate body temperature. structure and function of each type. In later chapters, we’ll
Many organs have more than one physiological role. If the discuss how these tissue types contribute to the functions of
roles are distinct enough, we consider the organ to belong to particular organ systems.
more than one organ system. The pancreas, for instance, pro-
duces enzymes critical to the function of the digestive system Mastering Biology Animation: Overview of Animal Tissues
Table 40.1 Organ Systems in Mammals
Organ System Main Components Main Functions

Digestive Mouth, pharynx, esophagus, stomach, intestines, liver, pancreas, Food processing (ingestion, digestion,
anus (See Figure 41.8.) absorption, elimination)
Circulatory Heart, blood vessels, blood (See Figure 42.5.) Internal distribution of materials
Respiratory Lungs, trachea, other breathing tubes (See Figure 42.24.) Gas exchange (uptake of oxygen; disposal
of carbon dioxide)
Immune and lymphatic Bone marrow, lymph nodes, thymus, spleen, lymph vessels Body defense (fighting infections and
(See Figure 43.6.) virally induced cancers)
Excretory Kidneys, ureters, urinary bladder, urethra (See Figure 44.12.) Disposal of metabolic wastes; regulation of
osmotic balance of blood
Endocrine Pituitary, thyroid, pancreas, adrenal, and other hormone- Coordination of body activities (such as
secreting glands (See Figure 45.8.) digestion and metabolism)
Reproductive Ovaries or testes and associated organs (See Figures 46.9 Gamete production; promotion of fertiliza-
and 46.10.) tion; support of developing embryo
Nervous Brain, spinal cord, nerves, sensory organs (See Figure 49.6.) Coordination of body activities; detection
of stimuli and formulation of responses
to them
Integumentary Skin and its derivatives (such as hair, claws, sweat glands) Protection against mechanical injury,
(See Figure 50.5.) infection, dehydration; thermoregulation
Skeletal Skeleton (bones, tendons, ligaments, cartilage) Body support, protection of internal
(See Figure 50.37.) organs, movement
Muscular Skeletal muscles (See Figure 50.26.) Locomotion and other movement

. Figure 40.5 Exploring Structure and Function in Animal Tissues
Epithelial Tissue
Occurring as sheets of cells, epithelial tissues, or epithelia (singu- Stratified squamous

lar, epithelium), cover the outside of the body and line organs and epithelium
cavities within the body. Because epithelial cells are closely packed,
often with tight junctions, they function as a barrier against Apical
mechanical injury, pathogens, and fluid loss. Epithelia also form surface
active interfaces with the environment. For example, the epithe-
lium that lines the nasal passages is crucial for Basal
olfaction, the sense of smell. Note surface
how different cell shapes and
arrangements correlate with
distinct functions. A stratified squamous epi-
thelium is multilayered and
Mastering Biology regenerates rapidly. New cells
Animation: Epithelial formed by division near the
Tissue basal surface push outward,
replacing cells that are sloughed
off. This epithelium is common-
ly found on surfaces subject to
abrasion, such as the outer skin
and the linings of the mouth,
anus, and vagina.
Cuboidal epithelium Simple columnar Simple squamous Pseudostratified

epithelium epithelium columnar epithelium
The single layer of platelike

A cuboidal epithelium, with cells that form a simple squa-
dice-shaped cells specialized mous epithelium functions in
The large, brick-shaped cells of A pseudostratified epithelium
for secretion, makes up the the exchange of material by dif-
simple columnar epithelia are consists of a single layer of cells
epithelium of kidney tubules fusion. This type of epithelium,
often found where secretion or varying in height and the posi-
and many glands, including which is thin and leaky, lines
active absorption is important. tion of their nuclei. In many
the thyroid gland and salivary blood vessels and the air sacs
For example, a simple columnar vertebrates, a pseudostratified
glands. of the lungs, where diffusion of
epithelium lines the intestines, epithelium of ciliated cells forms
nutrients and gases is essential.
secreting digestive juices and a mucous membrane that lines
absorbing nutrients. portions of the respiratory tract.
The beating cilia sweep the film
of mucus along the surface.
Lumen Apical surface

Polarity of epithelia
All epithelia are polarized, meaning that they have two differ-
ent sides. The apical surface faces the lumen (cavity) or outside
of the organ and is therefore exposed to fluid or air. Specialized
projections often cover this surface. For example, the apical sur-
face of the epithelium lining the small intestine is covered with
microvilli, projections that increase the surface area available
Basal surface for absorbing nutrients. Opposite the apical surface of each
epithelium is the basal surface.
10 om

. Figure 40.5 (continued) Exploring Structure and Function in Animal Tissues
Connective Tissue
Connective tissue, consisting of a sparse population of cells scat- reticular fibers join connective tissue to adjacent tissues, and
tered through an extracellular matrix, holds many tissues and elastic fibers make tissues elastic. If you pinch a fold of tissue
organs together and in place. The matrix generally on the back of your hand, the collagenous and reticular
consists of a web of fibers embedded in a liquid, fibers prevent the skin from being pulled far from the bone,
jellylike, or solid foundation. Within the matrix whereas the elastic fibers restore the skin to its original shape
are numerous cells called fibroblasts, which when you release your grip. Different mixtures of fibers
secrete fiber proteins, and macrophages, which and foundation form the major types of connective tissue
engulf foreign particles and any cell debris by shown below.
phagocytosis. Mastering Biology Animation:
Connective tissue fibers are of three kinds: Connective Tissue
Collagenous fibers provide strength and flexibility,
Blood
Blood has a liquid extracellular
matrix called plasma, which con-
Loose connective tissue sists of water, salts, and dissolved
proteins. Suspended in plasma
The most widespread connec- are erythrocytes (red blood cells),
Collagenous fiber
tive tissue in the vertebrate leukocytes (white blood cells),
body is loose connective tissue, and cell fragments called platelets.
which binds epithelia to Red cells carry oxygen, white cells
underlying tissues and holds function in defense, and platelets
organs in place. Loose con- aid in blood clotting.
nective tissue gets its name
120 om
from the loose weave of its Plasma

fibers, which include all three
types. It is found in the skin Elastic fiber White
and throughout the body. blood cells
50 om
Fibrous connective tissue
Fibrous connective tissue is
dense with collagenous fibers. Red blood cells
It is found in tendons, which
attach muscles to bones, and Adipose tissue Cartilage
in ligaments, which connect Adipose tissue is a specialized Cartilage contains collagenous
30 om
bones at joints. loose connective tissue that fibers embedded in a rubbery

stores fat in adipose cells dis- protein-carbohydrate complex
tributed throughout its matrix. called chondroitin sulfate.
Nuclei Adipose tissue pads and insu- Cells called chondrocytes secrete
lates the body and stores fuel as the collagen and chondroitin
fat molecules. Each adipose cell sulfate, which together make
Bone contains a large fat droplet that cartilage a strong yet flexible
The skeleton of most vertebrates is made of bone, swells when fat is stored and support material. The skeletons
a mineralized connective tissue. Bone-forming cells shrinks when the body uses that of many vertebrate embryos
called osteoblasts deposit a matrix of collagen. Calcium, fat as fuel. contain cartilage that is re-
magnesium, and phosphate ions combine into a hard placed by bone as the embryo
mineral within the matrix. The microscopic structure of matures. Cartilage remains in
hard mammalian bone consists of repeating units called Lipid droplets some locations, such as the
osteons. Each osteon has concentric layers of the min- disks that act as cushions
eralized matrix, which are deposited around a central between vertebrae.
150 om
canal containing blood vessels and nerves.
Central canal Chondrocytes

100 om
700 om
Osteon
Chondroitin sulfate

Muscle Tissue
The tissue responsible for nearly all types of body movement is muscle
tissue. All muscle cells consist of filaments containing the proteins
actin and myosin, which together enable muscles to contract. There
are three types of muscle tissue in the vertebrate body: skeletal,
smooth, and cardiac.
Mastering Biology Animation: Muscle Tissue
Skeletal muscle
Attached to bones by tendons, skeletal
muscle, or striated muscle, is responsible
for voluntary movements. Skeletal muscle
consists of bundles of long cells that are
Smooth muscle Cardiac muscle
called muscle fibers. During development,
skeletal muscle fibers form by the fusion of Smooth muscle, which lacks striations, Cardiac muscle forms the contractile
many cells, resulting in multiple nuclei in is found in the walls of the digestive wall of the heart. It is striated like skeletal
each muscle fiber. The arrangement of contract, urinary bladder, arteries, and other muscle and has similar contractile proper-
tractile units, or sarcomeres, along the internal organs. The cells are spindle- ties. Unlike skeletal muscle, however, car-
fibers gives the cells a striped (striated) shaped. Smooth muscles are responsible diac muscle has branched fibers that inter-
appearance. In adult mammals, build- for involuntary body activities, such as connect via intercalated disks, which relay
ing muscle increases the size but not the churning of the stomach and constric- signals from cell to cell and help synchronize
number of muscle fibers. tion of arteries. heart contraction.
Nuclei
Muscle
fiber
Sarcomere
100 om Nucleus Muscle fibers 25 om Nucleus Intercalated disk 25 om
Nervous Tissue
Nervous tissue functions in the receipt, processing, and transmission of information. Nervous tissue contains
neurons, or nerve cells, which transmit nerve impulses, as well as support cells called glial cells, or simply
glia. In many animals, a concentration of nervous tissue forms a brain, an information-processing center.
Mastering Biology Animation: Nervous Tissue
15 om
Neurons Glia
Glia
Neurons are the basic Neuron: The various types of glia
units of the nervous sys- Dendrites help nourish, insulate, and
tem. A neuron receives replenish neurons, and
nerve impulses from Cell body in some cases, modulate
other neurons via its neuron function.
cell body and multiple Axon
extensions called den-
drites. Neurons transmit
Axons of
impulses to neurons, neurons
45 om
muscles, or other cells

via extensions called
axons, which are often Blood
bundled together into (Fluorescent LM) vessel
nerves. (Confocal LM)

Coordination and Control . Figure 40.6 Signaling in the endocrine and nervous
systems.
For an animal’s tissues and organ systems to function
effectively, they must act in concert with one another. (a) Signaling by hormones (b) Signaling by neurons
For example, when the wolf shown in Figure 40.5 hunts,
blood flow is regulated to bring adequate nutrients and STIMULUS STIMULUS
gases to its leg muscles, which in turn are activated by the
brain in response to cues detected by the nose. What sig-
nals coordinate activity? How do the signals move within Endocrine
cell Cell body
the body?
of neuron
Animals have two major systems for coordinating and
controlling responses to stimuli: the endocrine and nervous Nerve Axon
systems (Figure 40.6). In the endocrine system, signal- impulse
Hormone
ing molecules released into the bloodstream by endocrine
cells are carried to all locations in the body. In the nervous Signal travels Signal travels
everywhere along axon to
system, neurons transmit signals along dedicated routes
via the a specific
connecting specific locations in the body. In each system, the bloodstream. location.
type of pathway used is the same regardless of whether the
signal’s ultimate target is at the other end of the body or just a
few cell diameters away.
The signaling molecules that are broadcast throughout Axon
Blood
the body by the endocrine system are called hormones. vessel Nerve
It takes seconds for hormones to be released into the blood- impulse
stream and carried throughout the body. The effects are often
long-lasting, however, because hormones can remain in the
bloodstream for minutes or even hours. Axon
Different hormones cause distinct effects, and only

cells that have receptors for a particular hormone respond
(Figure 40.6a). Depending on which cells have receptors for
that hormone, the hormone may have an effect in just a
single location or in sites throughout the body. For example,
thyroid-stimulating hormone (TSH) acts solely on cells in the
thyroid gland. They in turn release thyroid hormone, which Response: Limited to cells
Response: Limited to cells that connect by specialized
acts on nearly every body tissue to increase oxygen consump- that have a receptor ( ) for junctions to an axon that
tion and heat production. the signal transmits an impulse
In the nervous system, signals called nerve impulses travel
to specific target cells along communication lines consisting
mainly of axons (Figure 40.6b). Transmission in the nervous VISUAL SKILLS After comparing the two diagrams, explain why a
system is extremely fast; nerve impulses take only a frac- particular nerve impulse signal has only one physical pathway but a
particular hormone molecule can have multiple physical pathways.
tion of a second to reach the target and last only a fraction
of a second.
Nerve impulses can act on other neurons, on muscle cells, Because the two major communication systems of the
and on cells and glands that produce secretions. Unlike the body differ in signal type, transmission, speed, and dura-
endocrine system, the nervous system conveys information tion, it is not surprising that they are adapted to different
by the pathway the signal takes. For example, a person can functions. The endocrine system is especially well adapted
distinguish different musical notes because each note’s fre- for coordinating gradual changes that affect the entire body,
quency activates neurons in the ear that connect to slightly such as growth, development, reproduction, metabolic pro-
different locations in the brain. cesses, and digestion. The nervous system is well suited for
Communication in the nervous system usually involves directing immediate and rapid responses to the environment,
more than one type of signal. Nerve impulses travel along such as reflexes and other rapid movements. Nevertheless,
axons, sometimes over long distances, as changes in voltage. the two systems often work in close coordination. Both
In contrast, passing information from one neuron to another help maintain a stable internal environment, our next topic
often involves very short-range chemical signals. of discussion.

CONCEPT CHECK 40.1 fluid. In addition, conforming need not involve changes
in an internal variable: Many marine invertebrates, such
1. What properties do all types of epithelia share?
as spider crabs (genus Libinia), let their internal solute con-
2. VISUAL SKILLS Consider the idealized animal in Figure 40.4.
At which sites must oxygen cross a plasma membrane in centration conform to the relatively stable salinity of their
traveling from the external environment to the cytoplasm ocean environment.
of a body cell?
3. WHAT IF? Suppose you are standing at the edge of a cliff
and suddenly slip, barely managing to keep your balance and Homeostasis
avoid falling. As your heart races, you feel a burst of energy,
due in part to a surge of blood into dilated (widened) vessels The steady body temperature of a river otter and the
in your muscles and an upward spike in the level of glucose in stable concentration of solutes in a freshwater bass are
your blood. Why might you expect that this “fight-or-flight” examples of homeostasis, which means the maintenance
response requires both the nervous and endocrine systems?
of internal balance. In achieving homeostasis, animals
For suggested answers, see Appendix A.
maintain a “steady state”—a relatively constant inter-
nal environment—even when the external environment
changes significantly.
CONCEPT 40.2 Many animals exhibit homeostasis for a range of physi-
cal and chemical properties. For example, humans maintain
Feedback control maintains a fairly constant body temperature of about 37°C (98.6°F), a
the internal environment in blood pH within 0.1 pH unit of 7.4, and a blood glucose con-
centration that is predominantly in the range of 70–110 mg
many animals of glucose per 100 mL of blood.
Many organ systems play a role in managing an animal’s
internal environment, a task that can present a major chal-
lenge. Imagine if your body temperature soared every time
Mechanisms of Homeostasis
you took a hot shower or slurped a steaming bowl of soup. Homeostasis requires a control system. Before exploring
Faced with environmental fluctuations, animals manage their homeostasis in animals, let’s get a basic picture of how a
internal environment by either regulating or conforming. control system works by considering a nonliving example:
the regulation of room temperature. Let’s assume you
want to keep a room at 20°C (68°F), a comfortable tem-
Regulating and Conforming perature for normal activity. You set a control device—the
Compare the two sets of data in Figure 40.7. The river otter’s thermostat—to 20°C. A thermometer in the thermostat
body temperature is largely independent of that of the sur- monitors the room temperature. If the temperature falls
rounding water, whereas the largemouth
bass’s body warms or cools when the . Figure 40.7 The relationship between body and environmental temperatures in
water temperature changes. We can an aquatic temperature regulator and an aquatic temperature conformer. The river
convey these two trends by labeling otter regulates its body temperature, keeping it stable across a wide range of environmental
temperatures. The largemouth bass, meanwhile, allows its internal environment to conform
the otter a regulator and the bass a con- to the water temperature.
former with regard to body temperature.
An animal is a regulator for an envi- 40
ronmental variable if it uses internal River otter
mechanisms to control internal change (temperature regulator)
Body temperature (°C)
in the face of external fluctuation. In 30

contrast, an animal is a conformer if it
allows its internal condition to change Largemouth bass
in accordance with external changes in 20 (temperature
the particular variable. conformer)
An animal may allow some inter-
nal conditions to conform to the 10
environment but regulate others. For
instance, the bass conforms to the
temperature of the water in which it 0
0 10 20 30 40
lives, but regulates the solute concen-
Ambient (environmental) temperature (°C)
tration in its blood and interstitial

. Figure 40.8 A nonliving example of temperature that “damps” its stimulus (see Figure 1.10). This type of feed-
regulation: control of room temperature. Regulating room back regulation plays a major role in homeostasis in animals.
temperature depends on a sensor/control center (a thermostat)
For example, when you exercise vigorously, you produce heat,
that detects temperature change and activates mechanisms that
reverse that change. which increases your body temperature. Your nervous system
detects this increase and triggers sweating. The evaporation
of sweat from your skin then cools your body, helping return
body temperature to its set point and eliminating the stimulus.
Homeostasis is a dynamic equilibrium, an interplay
Thermostat turns between external factors that tend to change the internal
air conditioner on. environment and internal control mechanisms that oppose
Room temperature Room temperature
increases. decreases.
such changes. Note that physiological responses to stimuli
are not instantaneous, just as switching on a furnace does
not immediately warm a room. As a result, homeostasis
moderates but doesn’t eliminate changes in the internal
environment. Fluctuation is greater if a variable has a normal
ROOM TEMPERATURE
AT 20°C range—an upper and lower limit—rather than a set point.
(set point) This is equivalent to a heating system that is programmed
to produce heat when the room temperature drops to 19°C
(66°F) and to stop heating when the temperature reaches
21°C (70°F). Regardless of whether there is a set point or a
Room temperature Room temperature normal range, homeostasis is enhanced by adaptations that
increases. decreases. reduce fluctuations, such as insulation in the case of tempera-
Thermostat turns
heater on. ture and physiological buffers in the case of pH.
Unlike negative feedback, positive feedback is a control
mechanism that amplifies the stimulus. In animals, positive-
feedback loops do not play a major role in homeostasis, but
instead help drive processes to completion. During child-
DRAW IT Label at least one stimulus, response, and sensor/control
birth, for instance, the pressure of the baby’s head against
center in the above figure.
sensors near the opening of the mother’s uterus stimulates
below 20°C, the thermostat turns on a radiator, furnace, or the uterus to contract. These contractions result in greater
other heater (Figure 40.8). When the room exceeds 20°C, pressure against the opening of the uterus, heightening the
the thermostat switches off the heater. If the temperature contractions and thereby causing even greater pressure, ulti-
then drifts below 20°C, the thermostat activates another mately causing the baby to be born.
heating cycle. If the temperature instead rises above 20°C,
the thermostat activates a cooling mechanism, such as by Alterations in Homeostasis
turning on an air conditioner. The set points and normal ranges for homeostasis can change
Like a home heating system, the homeostatic control sys- under various circumstances. In fact, regulated changes in the
tem in animals maintains a variable, such as body tempera- internal environment are essential to normal body functions.
ture or solute concentration, at or near a particular value, or Some regulated changes occur during a particular stage in life,
set point. A fluctuation in the variable above or below the such as the radical shift in hormone balance that occurs dur-
set point serves as the stimulus detected by a sensor. The ing puberty. Other regulated changes are cyclic, such as the
sensor signals a control center, which triggers a response, a variation in hormone levels responsible for a woman’s men-
physiological activity that helps return the variable to the set strual cycle (see Figure 46.14).
point. In the home heating example, a drop in temperature In all animals (and plants), certain cyclic alterations in
below the set point acts as a stimulus, the thermostat serves metabolism reflect a circadian rhythm, a set of physiologi-
as the sensor and control center, and the heater produces cal changes that occur roughly every 24 hours (Figure 40.9).
the response. One way to observe this rhythm is to monitor body tem-
perature, which in humans typically undergoes a cyclic rise
Feedback Control in Homeostasis and fall of more than 0.6°C (1°F) in every 24-hour period.
If you examine the circuit in Figure 40.8, you can see that Remarkably, a biological clock maintains this rhythm even
either response (heating or cooling) reduces the stimulus (the when variations in human activity, room temperature, and
change in temperature) that triggered that response. The cir- light levels are minimized (see Figure 40.9a). A circadian
cuit thus displays negative feedback, a control mechanism rhythm is thus intrinsic to the body, although the biological

. Figure 40.9 Human circadian rhythm. . Figure 40.10 Acclimatization by mountain climbers in the
Himalayas. To lessen the risk of altitude sickness when ascending
Body Melatonin a high peak, climbers acclimatize by camping partway up the
Core body temperature (°C) temperature concentration mountain. Spending time at an intermediate altitude allows the
Melatonin concentration
37.1 60 circulatory and respiratory systems to become more efficient in
capturing and distributing oxygen at a lower concentration.
in blood (pg/mL)
36.9 40
36.7 20
36.5 0
2 6 10 2 6 10
PM PM PM AM AM AM
Time of day
(a) Variation in core body temperature and melatonin concentra-

tion in blood. Researchers studied resting but awake volunteers in
an isolation chamber with constant temperature and low light.
(Melatonin is a hormone secreted by the pineal gland.)
Midnight
Start of Lowest
melatonin secretion heart rate
SLE
Greatest EP
Lowest body
muscle strength
temperature
6 PM 6 AM
Homeostasis is sometimes altered by acclimatization,
Most rapid
rise in blood an animal’s physiological adjustment to changes in its
pressure external environment. For instance, when an elk moves
Fastest
reaction time up into the mountains from sea level, the lower oxygen
Highest risk concentration in the high mountain air stimulates the
Noon of cardiac arrest animal to breathe more rapidly and deeply. As a result,
more CO2 is lost through exhalation, raising blood pH
(b) The human circadian clock. Metabolic activities undergo daily above its normal range. As the animal acclimatizes over
cycles in response to the circadian clock. As illustrated for a typical several days, changes in kidney function cause it to
individual who rises early in the morning, eats lunch around noon,
and sleeps at night, these cyclic changes occur throughout a excrete urine that is more alkaline, returning blood pH
24-hour day. to its normal range. Other mammals, including humans,
are also capable of acclimatizing to dramatic altitude
changes (Figure 40.10), although health risks remain.
clock is normally coordinated with the cycle of light and

darkness in the environment (see Figure 40.9b). For example, CONCEPT CHECK 40.2
the hormone melatonin is secreted at night, and more is 1. MAKE CONNECTIONS How does negative feedback in
released during the longer nights of winter. External stimuli thermoregulation differ from feedback inhibition in an
can reset the biological clock, but the effect is not immediate. enzyme-catalyzed biosynthetic process (see Figure 8.21)?
That is why flying across several time zones results in jet lag, 2. If you were deciding where to put the thermostat in a
house, what factors would govern your decision? How do
a mismatch between the circadian rhythm and local environ- these factors relate to the fact that many homeostatic con-
ment that persists until the clock fully resets. trol sensors in humans are located in the brain?
Noting the importance of biological clocks to human 3. MAKE CONNECTIONS Like animals, cyanobacteria have a
health and disease, the Nobel Prize Committee awarded the circadian rhythm. By analyzing the genes that maintain bio-
logical clocks, scientists concluded that the 24-hour rhythms
2017 Nobel Prize in Physiology or Medicine to Americans
of humans and cyanobacteria reflect convergent evolution
Jeffrey Hall, Michael Rosbash, and Michael Young, who stud- (see Concept 26.2). What evidence would have supported
ied the fruit fly Drosophila to map out the molecular mecha- this conclusion? Explain.
nisms that underlie circadian rhythms. For suggested answers, see Appendix A.

CONCEPT 40.3 Variation in Body Temperature
Animals also differ in whether their body temperature is vari-
Homeostatic processes for able or constant. An animal whose body temperature varies
thermoregulation involve form, with its environment is called a poikilotherm (from the Greek
poikilos, varied). In contrast, a homeotherm has a relatively
function, and behavior constant body temperature. For example, the largemouth
In this section, we’ll examine the regulation of body tempera- bass is a poikilotherm, and the river otter is a homeotherm
ture as an example of how form and function work together (see Figure 40.7).
in regulating an animal’s internal environment. Later chap- From the descriptions of ectotherms and endotherms, it
ters in this unit will discuss other physiological systems might seem that all ectotherms are poikilothermic and all
involved in maintaining homeostasis. endotherms are homeothermic. In fact, there is no fixed rela-
Thermoregulation is the process by which animals tionship between the source of heat and the stability of body
maintain their body temperature within a normal range. temperature. Many ectothermic marine fishes and inver-
Body temperatures outside the normal range can reduce the tebrates inhabit waters with such stable temperatures that
efficiency of enzymatic reactions, alter the fluidity of cellular their body temperature varies less than that of mammals and
membranes, and affect other temperature-sensitive biochemi- other endotherms. Conversely, the body temperature of a few
cal processes, potentially with fatal results. endotherms varies considerably. For example, the body tem-
In talking about thermoregulation, we will need to perature of some bats drops from 40°C to a few degrees above
talk about heat. Formally, heat is defined as thermal zero when they enter hibernation.
energy in transfer from one body of matter to another (see
Concept 8.1). Here, however, we will use the term heat to
refer simply to thermal energy. . Figure 40.11 Thermoregulation by internal or external
sources of heat. Endotherms obtain heat from their internal
metabolism, whereas ectotherms rely on heat from their
Endothermy and Ectothermy external environment.
Heat for thermoregulation can come from either internal
metabolism or the external environment. Humans and other
mammals, as well as birds, are endothermic, meaning that
they are warmed mostly by heat generated by metabolism.
Some fishes and insect species and a few nonavian reptiles
are also mainly endothermic. In contrast, amphibians,
many nonavian reptiles and fishes, and most invertebrates
are ectothermic, meaning that they gain most of their heat
from external sources. Endothermy and ectothermy are not
mutually exclusive, however. For example, a bird is mainly
endothermic but may warm itself in the sun on a cold morn-
ing, much as an ectothermic lizard does.
Endotherms can maintain a stable body temperature even
(a) King penguins (Aptenodytes patagonicus), endotherms
in the face of large fluctuations in the environmental tem-
perature. In a cold environment, an endotherm generates
enough heat to keep its body substantially warmer than its
surroundings (Figure 40.11a). In a hot environment, endo-
thermic vertebrates have mechanisms for cooling their bod-
ies, enabling them to withstand temperatures that are intoler-
able for most ectotherms.
Many ectotherms adjust their body temperature
by behavioral means, such as seeking out shade or bask-
ing in the sun (Figure 40.11b). Because their heat source
is largely environmental, ectotherms generally need to
consume much less food than endotherms of equiva-
lent size—an advantage if food supplies are limited.
Ectotherms also usually tolerate larger fluctuations in
their internal temperature. (b) Florida red-bellied turtles (Pseudemys nelsoni ), ectotherms

It is a common misconception that ectotherms are “cold- outer covering of the body, consisting of the skin, hair, and nails
blooded” and endotherms are “warm-blooded.” Ectotherms (claws or hooves in some species).
do not necessarily have low body temperatures. On the con-
trary, when sitting in the sun, many ectothermic lizards have Insulation
higher body temperatures than mammals. Thus, the terms Insulation, which reduces the flow of heat between an animal’s
cold-blooded and warm-blooded are misleading and are avoided body and its environment, is a major adaptation for thermo-
in scientific communication. regulation in both mammals and birds. Insulation is found
both at the body surface—hair and feathers—and beneath—
layers of fat formed by adipose tissue. In addition, some ani-
Balancing Heat Loss and Gain mals secrete oily substances that repel water, protecting the
Thermoregulation depends on an animal’s ability to control insulating capacity of feathers or fur. Birds, for example, secrete
the exchange of heat with its environment. That exchange oils that they apply to their feathers during preening.
can occur by any of four processes: radiation, evaporation, Often, animals can adjust their insulating layers to further
convection, and conduction (Figure 40.12). In each, heat is regulate body temperature. Most land mammals and birds,
transferred from an object of higher temperature to one of lower for example, react to cold by raising their fur or feathers. This
temperature. action traps a thicker layer of air, thereby increasing the effec-
The essence of thermoregulation is maintaining a rate of heat tiveness of the insulation. Lacking feathers or fur, humans
gain that equals the rate of heat loss. Animals do this through must rely primarily on fat for insulation. We do, however, get
mechanisms that either reduce heat exchange overall or favor “goose bumps,” a vestige of hair raising inherited from our
heat exchange in a particular direction. In mammals, several of furry ancestors.
these mechanisms involve the integumentary system, the Insulation is particularly important for marine mammals,
such as whales and walruses. These animals swim in water
. Figure 40.12 Heat exchange between an organism and its colder than their body core, and many species spend at least
environment. part of the year in nearly freezing polar seas. Furthermore, the
Radiation is the emission of Evaporation is the removal of transfer of heat to water occurs 50 to 100 times more rapidly
electromagnetic waves by all heat from the surface of a than heat transfer to air. Survival under these conditions is
objects warmer than absolute liquid that is losing some of its
zero. Here, a lizard absorbs molecules as gas. Evaporation made possible by an evolutionary adaptation called blubber,
heat radiating from the distant of water from a lizard‘s moist a very thick layer of insulating fat just under the skin. The
sun and radiates a smaller surfaces that are exposed to insulation that blubber provides is so effective that marine
amount of energy to the the environment has a strong
surrounding air. cooling effect. mammals can maintain body core temperatures of about
36938°C 1979100°F2 without requiring much more energy
from food than land mammals of similar size.
Circulatory Adaptations
Circulatory systems provide a major route for heat flow
between the interior and exterior of the body. Adaptations
that regulate the extent of blood flow near the body surface
or that trap heat within the body core play a significant role
in thermoregulation.
In response to changes in the temperature of their sur-
roundings, many animals alter the amount of blood (and
hence heat) flowing between their body core and their
skin. Nerve signals that relax the muscles of the vessel walls
result in vasodilation, a widening of superficial blood ves-
sels (those near the body surface). As a consequence of the
Convection is the transfer of Conduction is the transfer increase in vessel diameter, blood flow in the skin increases.
heat by the movement of air or of heat between molecules
liquid past a surface, as when a of objects in contact with In endotherms, vasodilation usually increases the transfer
breeze contributes to heat loss each other, as when a of body heat to the environment by radiation, conduc-
from a lizard‘s dry skin or when lizard sits on a hot rock. tion, and convection (see Figure 40.12). The reverse process,
blood moves heat from the
body core to the extremities. vasoconstriction, reduces blood flow and heat transfer by
decreasing the diameter of superficial vessels.
VISUAL SKILLS If this figure showed a penguin (an endotherm) on an
ice floe rather than an iguana (an ectotherm) on a rock, would any of the Like endotherms, some ectotherms control heat exchange
arrows point in a different direction? Explain. by regulating blood flow. For example, when the marine

. Figure 40.13 Countercurrent heat exchangers. A countercurrent exchange system traps heat in
the body core, thus reducing heat loss from the extremities, particularly when they are immersed in cold
water or in contact with ice or snow. In essence, heat in the arterial blood emerging from the body core
is transferred directly to the returning venous blood instead of being lost to the environment.
Canada goose 1
tates heat transfer from arteries to

veins along the entire length of
the blood vessels.
2 1
1 3 Vein
Artery Vein
Artery
can still transfer heat to the even colder 3
358C 338
blood in an adjacent vein. The blood in the 3
308 278
Arteries carry
blood into the 208 188 3
2
foot. Veins
108 98
to the heart.
Key
2
Warm blood
arteries and veins.
To From Cool blood Heat transfer
foot foot
iguana of the Galápagos Islands swims in the cold ocean, endothermic insects (bumblebees, honeybees, and some
its superficial blood vessels undergo vasoconstriction. moths) have a countercurrent exchanger that helps
This process routes more blood to the body core, conserving maintain a high temperature in their thorax, where flight
body heat. muscles are located.
In many birds and mammals, reducing heat loss
from the body relies on countercurrent exchange,
the transfer of heat (or solutes) between fluids that are Cooling by Evaporative Heat Loss
flowing in opposite directions. In a countercurrent heat Many mammals and birds live in places where regulat-
exchanger, arteries and veins are located adjacent to each ing body temperature requires cooling at some times
other (Figure 40.13). Because blood flows through the and warming at others. If environmental temperature
arteries and veins in opposite directions, this arrangement is above body temperature, only evaporation can keep
allows heat exchange to be remarkably efficient. As warm body temperature from rising. Water absorbs consider-
blood in the arteries moves outward from the body core, able heat when it evaporates (see Concept 3.2); this heat
it transfers heat to the colder blood in the veins returning is carried away from the skin and respiratory surfaces with
from the extremities. Most importantly, heat is transferred water vapor.
along the entire length of the exchanger, maximizing Some animals exhibit adaptations that greatly facilitate
the rate of heat exchange and minimizing heat loss to evaporative cooling. A few mammals, including horses and
the environment. humans, have sweat glands. In many other mammals, as well
Although most sharks and fishes are temperature as in birds, panting is important. Some birds have a pouch
conformers, countercurrent heat exchangers are found richly supplied with blood vessels in the floor of the mouth;
in some large, powerful swimmers, including great fluttering the pouch increases evaporation. Pigeons can
white sharks, bluefin tuna, and swordfish. By keep- use this adaptation to keep their body temperature close to
ing the main swimming muscles warm, this adaptation 40°C (104°F) in air temperatures as high as 60°C (140°F), as
enables vigorous, sustained activity. Similarly, many long as they have sufficient water.

c Figure 40.14 chickadees, birds with a body mass of only 20 g, remain
Thermoregulatory behavior active and hold their body temperature nearly constant
in a dragonfly. By orienting its
at 40°C (104°F) in environmental temperatures as low as
body so that the narrow tip of
its abdomen faces the sun, the - 40°C 1- 40°F2.
dragonfly minimizes heating Flying insects such as bees and moths can also vary
by solar radiation. heat production. Many such endothermic insects warm
up by shivering before taking off. As they contract their flight
muscles in synchrony, only slight wing movements occur,
Behavioral Responses but considerable heat is produced. Chemical reactions, and
Ectotherms, and hence cellular respiration, accelerate in the warmed-up flight
sometimes endotherms, “motors,” enabling flight even in cold air.
control body tempera- In some mammals, endocrine signals released in response
ture through behavioral to cold cause mitochondria to increase their metabolic activity
responses to changes and produce heat instead of ATP. This process, called nonshiv-
in the environment. ering thermogenesis, takes place throughout the body. Some
When cold, they seek mammals also have a tissue called brown fat in their neck and
warm places, orienting between their shoulders that is specialized for rapid heat pro-
themselves toward heat duction. (The presence of extra mitochondria is what gives
sources and expanding brown fat its characteristic color.) Brown fat is found in the
the portion of their body surface exposed to the heat source infants of many mammals, representing about 5% of total body
(see Figure 40.11b). When hot, they bathe, move to cool weight in human infants. Long known to be present in adult
areas, or turn in another direction, minimizing their absorp- mammals that hibernate, brown fat has also recently been
tion of heat from the sun. For example, a dragonfly’s “obe- detected in human adults (Figure 40.15). There, the amount
lisk” posture is an adaptation that minimizes the amount has been found to vary, with individuals exposed to a cool envi-
of body surface exposed to the sun and thus to heating ronment for a month having increased amounts of brown fat.
(Figure 40.14). Although these behaviors are relatively sim- Among the nonavian reptiles, endothermy has been
ple, they enable many ectotherms to maintain a nearly con- observed in some large species in certain circumstances. For
stant body temperature. example, researchers found that a female Burmese python
Social behavior contributes to thermoregulation in both (Python molurus bivittatus) incubating eggs maintained a
endotherms and ectotherms. Among endotherms, for body temperature roughly 6°C (11°F) above that of the sur-
example, behavior contributes significantly to the winter rounding air. Where did the heat come from? Further studies
survival of Emperor penguins (see Figure 40.1). Among showed that such pythons, like birds, can raise their body
ectotherms, honeybees are notable for their use of behavior
in achieving homeostasis for temperature. In cold weather,
they increase heat production and huddle together, thereby . Figure 40.15 Brown fat activity during cold stress. This
PET scan shows metabolically active brown fat deposits (indicated
retaining heat. Individuals move between the cooler by the arrows) surrounding the neck.
outer edges of the huddle and the warmer center, thus
circulating and distributing the heat. In hot weather,
honeybees cool the hive by transporting water to the
hive and fanning with their wings, promoting evapora-
tion and convection. Thus, a honeybee colony uses many
of the mechanisms of thermoregulation characteristic of
individual animals.
Adjusting Metabolic Heat Production

Because endotherms generally maintain a body temperature
considerably higher than that of the environment, they
must counteract continual heat loss. Endotherms can vary
heat production—thermogenesis—to match changing rates of
heat loss. Thermogenesis is increased by such muscle activ-
ity as moving or shivering. For example, shivering helps

▼ Figure 40.16 Inquiry to seasonal temperature changes often includes adjusting
insulation—growing a thicker coat of fur in the winter and
How does a Burmese python generate heat shedding it in the summer, for example.
while incubating eggs? Acclimatization in ectotherms often includes adjust-
ments at the cellular level. Cells may produce variants of
Experiment Herndon Dowling and colleagues at the Bronx
Zoo in New York observed that when a female Burmese py- enzymes that have the same function but different opti-
thon incubated eggs by wrapping her body around them, mal temperatures. Also, the proportions of saturated and
she raised her body temperature and frequently contracted unsaturated lipids in membranes may change; unsaturated
the muscles in her coils. To learn if the contractions were
lipids help keep membranes fluid at lower temperatures
elevating her body temperature, they placed the python and
her eggs in a chamber. As they varied the chamber’s temper- (see Figure 7.5).
ature, they monitored the python’s muscle contractions as Remarkably, some ectotherms can survive subzero
well as her oxygen uptake, a measure of her rate of cellular temperatures, producing “antifreeze” proteins that
respiration.
prevent ice formation in their cells. In the Arctic and
Results The python’s oxygen consumption increased when Southern (Antarctic) Oceans, these proteins enable cer-
the temperature in the chamber decreased. As shown in the
tain fishes to survive in water as cold as - 2°C 128°F2, a full
graph, this increase in oxygen consumption paralleled an
increase in the rate of muscle contraction. degree Celsius below the freezing point of body fluids in
other species.
120
Physiological Thermostats and Fever
O2 consumption (mL O2/hr • kg)
100
In humans and other mammals, the sensors responsible for
80 thermoregulation are concentrated in the hypothalamus,
the brain region that also controls the circadian clock. Within
60 the hypothalamus, a group of nerve cells functions as a ther-
mostat, responding to body temperatures outside the normal
40 range by activating mechanisms that promote heat loss or
gain (Figure 40.17).
20
At body temperatures above the normal range, the hypo-
0 thalamic thermostat promotes cooling of the body
0 5 10 15 20 25 30 35 by dilation of vessels in the skin, sweating, or panting. When
Contractions per minute body temperatures instead drop below the normal range,
the thermostat inhibits heat loss mechanisms and activates
Conclusion Because oxygen consumption, which gen- mechanisms that either save heat, such as constricting
erates heat through cellular respiration, was correlated vessels in the skin, or generate heat, such as shivering.
with the rate of muscle contraction, the researchers con-
cluded that the muscle contractions, a form of shivering, In the course of certain bacterial and viral infections,
were the source of the Burmese python’s elevated body mammals and birds develop fever, an elevated body tem-
temperature. perature. A variety of experiments have shown that fever
Data from V. H. Hutchison, H. G. Dowling, and A. Vinegar, Thermoregulation reflects an increase in the normal range for the biological
in a brooding female Indian python, Python molurus bivittatus, Science
thermostat. For example, artificially raising the temperature
151:694–696 (1966).
of the hypothalamus in an infected animal reduces fever in
WHAT IF? Suppose you varied air temperature and measured
oxygen consumption for a female Burmese python without a clutch of the rest of the body.
eggs. Since she would not show shivering behavior, how would you Among certain ectotherms, an increase in body tem-
expect the snake’s oxygen consumption to vary with environmental perature upon infection reflects what is called a behavioral
temperature?
fever. For example, the desert iguana (Dipsosaurus dorsalis)
responds to infection with certain bacteria by seeking
a warmer environment and then maintaining a body
temperature through shivering (Figure 40.16). Whether cer- temperature that is elevated by 2–4°C (4–7°F). Similar obser-
tain groups of Mesozoic dinosaurs were similarly endother- vations in fishes, amphibians, and even cockroaches indicate
mic is a matter of active debate. that fever is common to both endotherms and ectotherms.
Now that we have explored thermoregulation, we’ll
Acclimatization in Thermoregulation conclude our introduction to animal form and function by
Acclimatization contributes to thermoregulation in many considering the different ways that animals allocate, use,
animal species. In birds and mammals, acclimatization and conserve energy.

c Figure 40.17 The thermostatic Blood vessels in skin
function of the hypothalamus in dilate; capillaries fill
human thermoregulation. with warm blood; heat
Thermostat in radiates from skin surface.
WHAT IF? Suppose at the end of a hard run hypothalamus
on a hot day you find that there are no drinks activates cooling
left in the cooler. If, out of desperation, you mechanisms.
dunk your head into the cooler, how might Sweat glands
the ice-cold water affect the rate at which secrete
your body temperature returns to normal? sweat, which
evaporates,
Mastering Biology Figure Walkthrough cooling the
Body temperature increases
body. Body temperature
(such as when exercising or
in hot surroundings). decreases.
NORMAL BODY TEMPERATURE

(approximately 36–38°C)
Body temperature decreases

Body temperature (such as when in cold
increases. surroundings).
Skeletal muscles rapidly
contract, causing shivering,
which generates heat.
Blood vessels in skin Thermostat in

constrict, diverting blood hypothalamus
from skin to deeper activates warming
tissues and reducing heat mechanisms.
loss from skin surface.
CONCEPT CHECK 40.3 reproduction. The overall flow and transformation of

1. What mode of heat exchange is involved in “wind chill,” energy in an animal—its bioenergetics—determines
when moving air feels colder than still air at the same nutritional needs and is related to the animal’s size, activ-
temperature? Explain. ity, and environment.
2. Flowers differ in how much sunlight they absorb. Why
might this matter to a hummingbird seeking nectar on a
cool morning? Energy Allocation and Use
3. WHAT IF? Why is shivering likely during the onset of a fever? Organisms can be classified by how they obtain chemical
For suggested answers, see Appendix A. energy. Most autotrophs, such as plants, harness light energy
to build energy-rich organic molecules and then use those
molecules for fuel. Most heterotrophs, such as animals, obtain
CONCEPT 40.4 their chemical energy from food, which contains organic
molecules synthesized by other organisms.
Energy requirements are related Animals use chemical energy harvested from the food they
to animal size, activity, and eat to fuel metabolism and activity. Food is digested by enzy-
matic hydrolysis (see Figure 5.2b), and nutrients are absorbed
environment by body cells. The ATP (adenosine triphosphate) produced by
One of the unifying themes of biology, introduced cellular respiration and fermentation powers cellular work,
in Concept 1.1, is that life requires energy transfer enabling cells, organs, and organ systems to perform the
and transformation. Like other organisms, animals functions that keep an animal alive. Other uses of energy in
use chemical energy for growth, repair, activity, and the form of ATP include biosynthesis, which is needed for

. Figure 40.18 Bioenergetics of an animal: an overview. kilocalories (kcal). A kilocalorie equals 1,000 calories, or
4,184 joules. (The unit Calorie, with a capital C, as used by
Organic molecules many nutritionists, is actually a kilocalorie.)
in food Metabolic rate can be determined in several ways.
External
environment Because nearly all of the chemical energy used in cellular respi-
Animal ration eventually appears as heat, metabolic rate can be mea-
body Digestion and sured by monitoring an animal’s rate of heat loss. For this
absorption Heat
approach, researchers use a calorimeter, which is a closed,
Energy
insulated chamber equipped with a device that records the
lost in
feces heat an animal gives off to its environment. Metabolic rate
Nutrient molecules can also be determined from the amount of oxygen consumed
in body cells Energy
lost in or carbon dioxide produced by an animal’s cellular respira-
nitrogenous tion (Figure 40.19). To calculate metabolic rate over longer
waste
periods, researchers record the rate of food consumption,
Carbon Cellular the energy content of the food (about 4.5–5 kcal per gram of
Heat
skeletons respiration protein or carbohydrate and about 9 kcal per gram of fat), and
the chemical energy lost in waste products (feces and urine or
other nitrogenous wastes).
ATP
Biosynthesis:
growth, Minimum Metabolic Rate
storage, and
reproduction and Thermoregulation
Cellular Heat Animals must maintain a minimum metabolic rate for basic
work
functions such as cell maintenance, breathing, and circula-
tion. Researchers measure this minimum metabolic rate
Heat differently for endotherms and ectotherms. The minimum
metabolic rate of a nongrowing endotherm that is at rest,
MAKE CONNECTIONS Use the idea of energy coupling to
explain why heat is produced in the absorption of nutrients, has an empty stomach, and is not experiencing stress is
in cellular respiration, and in the synthesis of biopolymers called the basal metabolic rate (BMR). BMR is measured
(see Concept 8.3).
under a “comfortable” temperature range—a range that
requires only the minimum generation or shedding of heat.
The minimum metabolic rate of ectotherms is determined
body growth and repair, synthesis of storage material such at a specific temperature because changes in the environ-
as fat, and production of gametes. The production and use of mental temperature alter body temperature and therefore
ATP generate heat, which the animal eventually gives off to metabolic rate. The metabolic rate of a fasting, nonstressed
its surroundings (Figure 40.18).
. Figure 40.19 Measuring the rate of oxygen consumption by a swimming shark.

Quantifying Energy Use A researcher monitors the decrease in oxygen level over time in the recirculating water of a
juvenile hammerhead’s tank.
How much of the total energy an ani-
mal obtains from food does it need just
to stay alive? How much energy must
be expended to walk, run, swim, or fly
from one place to another? What fraction
of the energy intake is used for reproduc-
tion? Physiologists answer such questions
by measuring the rate at which an animal
uses chemical energy and how this rate
changes in different circumstances.
The sum of all the energy an animal
uses in a given time interval is called its
metabolic rate. Energy is measured
in joules (J) or in calories (cal) and

ectotherm at rest at a particular temperature is called its . Figure 40.20 The relationship of metabolic rate to body size.
standard metabolic rate (SMR).
Comparisons of minimum metabolic rates reveal the 103
different energy costs of endothermy and ectothermy. Elephant
The BMR for humans averages 1,600–1,800 kcal per day 102 Horse
BMR (L O2/hr) (log scale)

for adult males and 1,300–1,500 kcal per day for adult
Human
females. These BMRs are about equivalent to the rate of
10 Sheep
energy use by a 75-watt lightbulb. In contrast, the SMR of an
American alligator is only about 60 kcal per day at Dog
Cat
20°C (68°F). As this represents less than 1⁄20 the energy 1
used by a comparably sized adult human, it is clear that
ectothermy has a markedly lower energetic requirement Rat
than endothermy. 10–1 Ground squirrel
Shrew
Mouse
Harvest mouse
10–2
Influences on Metabolic Rate 10–3 10–2 10–1 1 10 102 103
Metabolic rate is affected by many factors other than an ani- Body mass (kg) (log scale)
mal being an endotherm or an ectotherm. Some key factors
(a) Relationship of basal metabolic rate (BMR) to body size for various
are age, sex, size, activity, temperature, and nutrition. Here
mammals. From shrew to elephant, size increases 1 millionfold.
we’ll examine the effects of size and activity.
8
Size and Metabolic Rate Shrew
BMR per kg body mass (L O2/hr • kg)

Larger animals have more body mass and therefore require 7
more chemical energy. Remarkably, the relationship between 6
overall metabolic rate and body mass is constant across a
wide range of sizes and forms, as illustrated for various mam- 5
mals in Figure 40.20a. In fact, for even more varied organisms 4
ranging in size from bacteria to blue whales, metabolic rate
3 Harvest mouse
remains roughly proportional to body mass to the three-
quarter power (m3/4). Scientists are still researching the basis 2 Mouse Sheep
of this relationship, which applies to ectotherms as well Rat
Cat Human Elephant
as endotherms. 1 Dog
Ground squirrel Horse
The relationship of metabolic rate to size profoundly 0
affects energy consumption by body cells and tissues. 10–3 10–2 10–1 1 10 102 103
As shown in Figure 40.20b, the energy it takes to main- Body mass (kg) (log scale)
tain each gram of body mass is inversely related to body
(b) Relationship of BMR per kilogram of body mass to body size for
size. Each gram of a mouse, for instance, requires about
the same mammals as in (a).
20 times as many calories as a gram of an elephant, even
though the whole elephant uses far more calories than the INTERPRET THE DATA Based on the graph in (a), one observer
whole mouse. The smaller animal’s higher metabolic rate suggests that a group of 100 ground squirrels has the same basal
metabolic rate as 1 dog. A second observer looking at the graph
per gram demands a higher rate of oxygen delivery. To meet disagrees. Who is correct and why?
this demand, the smaller animal must have a higher breath-
ing rate, blood volume (relative to its size), and heart rate.
Thinking about body size in bioenergetic terms reveals insect twitching its wings consumes energy beyond the BMR
how trade-offs shape the evolution of body plans. As body or SMR. Maximum metabolic rates (the highest rates of ATP
size decreases, each gram of tissue increases in energy cost. use) occur during peak activity, such as lifting a heavy object,
As body size increases, energy costs per gram of tissue sprinting, or swimming at high speed. In general, the maxi-
decrease, but an ever-larger fraction of body tissue is required mum metabolic rate an animal can sustain is inversely related
for exchange, support, and locomotion. to the duration of activity.
For most terrestrial animals, the average daily rate of
Activity and Metabolic Rate energy consumption is two to four times BMR (for endo-
For both ectotherms and endotherms, activity greatly affects therms) or SMR (for ectotherms). Humans in most developed
metabolic rate. Even a person reading quietly at a desk or an countries have an unusually low average daily metabolic rate

Scientific Skills Exercise
Interpreting Pie Charts
How Do Energy Budgets Differ for Three Terrestrial
Vertebrates? To explore bioenergetics in animal bodies,
let’s consider typical annual energy budgets for three terrestrial
vertebrates that vary in size and thermoregulatory strategy:
a 4-kg male Adélie penguin, a 25-g (0.025-kg) female deer
mouse, and a 4-kg female ball python. The penguin is well-
insulated against his Antarctic environment but must expend
energy in swimming to catch food, incubating eggs laid by his
partner, and bringing food to his chicks. The tiny deer mouse
lives in a temperate environment where food may be read-
ily available, but her small size causes rapid loss of body heat.
Unlike the penguin and mouse, the python is ectothermic and Adélie penguin Deer mouse Ball python
keeps growing throughout her life. She produces eggs but does 4-kg male 0.025-kg female 4-kg female
not incubate them. In this exercise, we’ll compare the energy 340,000 kcal/yr 4,000 kcal/yr 8,000 kcal/yr
expenditures of these animals for five important functions:
Key
basal (standard) metabolism, reproduction, thermoregulation,
activity, and growth. Basal (standard) metabolism Activity
Reproduction Growth
How the Data Were Obtained Energy budgets were calculated
for each of the animals based on measurements from field and Thermoregulation
laboratory studies.
Data from M. A. Chappell et al., Energetics of foraging in breeding Adélie pen-
guins, Ecology 74:2450–2461 (1993); M. A. Chappell et al., Voluntary running
Data from the Experiments Pie charts are a good way to in deer mice: speed, distance, energy costs, and temperature effects, Journal
compare relative differences in a set of variables. In the pie of Experimental Biology 207:3839–3854 (2004); T. M. Ellis and M. A. Chappell,
charts here, the sizes of the wedges represent the relative Metabolism, temperature relations, maternal behavior, and reproductive en-
annual energy expenditures for the functions shown in the ergetics in the ball python (Python regius), Journal of Comparative Physiology B
157:393–402 (1987).
key. The total annual expenditure for each animal is given
below its pie chart. 4. Now look at the total annual energy expenditures for each ani-
mal. How much more energy does the penguin expend each year
INTERPRET THE DATA
compared to the similarly sized python?
1. You can estimate the contribution of each wedge in a pie
5. Which animal expends the most kilocalories per year on
chart by remembering that the entire circle represents 100%,
thermoregulation?
half is 50%, and so on. What percent of the mouse’s energy
budget goes to basal metabolism? What percent of the 6. If you monitored energy allocation in the penguin for just
penguin’s budget is for activity? a few months instead of an entire year, you might find the
growth category to be a significant part of the pie chart. Given
2. Without considering the sizes of the wedges, how do the three
that adult penguins don’t grow from year to year, how would
pie charts differ in which functions they include? Explain these
you explain this finding?
differences.
3. Does the penguin or the mouse expend a greater proportion Instructors: A version of this Scientific Skills Exercise can be
of its energy budget on thermoregulation? Why? assigned in Mastering Biology.
of about 1.5 times BMR—an indication of a relatively seden- A major adaptation that enables animals to save energy in the
tary lifestyle. face of such difficult conditions is torpor, a physiological
The fraction of an animal’s energy “budget” that is state of decreased activity and metabolism.
devoted to activity depends on many factors, including its Many birds and small mammals exhibit a daily torpor that
environment, behavior, size, and thermoregulation. In the is well adapted to feeding patterns. For instance, some bats
Scientific Skills Exercise, you’ll interpret data on the annual feed at night and go into torpor in daylight. Similarly, chicka-
energy budgets of three terrestrial vertebrates. dees and hummingbirds, which feed during the day, often go
into torpor on cold nights.
All endotherms that exhibit daily torpor are relatively
Torpor and Energy Conservation small; when active, they have high metabolic rates and thus
Despite their many adaptations for homeostasis, animals may very high rates of energy consumption. The changes in body
encounter conditions that severely challenge their abilities to temperature, and thus the energy savings, are often consid-
balance their heat, energy, and materials budgets. For exam- erable: the body temperature of chickadees drops as much
ple, at certain times of the day or year, their surroundings as 10°C (18°F) at night, and the core body temperature of a
may be extremely hot or cold, or food may be unavailable. hummingbird can fall 25°C (45°F) or more.

Hibernation is long- . Figure 40.21 A hazel ▼ Figure 40.22 Inquiry
term torpor that is an adap- dormouse (Muscardinus
avellanarius) hibernating. What happens to the circadian clock during
tation to winter cold and
food scarcity. When a mam- hibernation?
mal enters hibernation, its
Experiment To determine whether the 24-hour biological
body temperature declines clock continues to run during hibernation, Paul Pévet and
as its body’s thermostat is colleagues at the University of Louis Pasteur in Strasbourg,
turned down (Figure 40.21). France, studied molecular components of the circadian
clock in the European hamster (Cricetus cricetus). The
Some hibernating mam-
researchers measured RNA levels for two clock genes—
mals cool to as low as Per2 and Bmal1—during normal activity (euthermia) and
192°C 134936°F2, during hibernation in constant darkness. The RNA samples
and at least one, the were obtained from the suprachiasmatic nuclei (SCN), a
pair of structures in the mammalian brain that control
Arctic ground squirrel
circadian rhythms.
(Spermophilus parryii), can enter a supercooled (unfrozen)
state in which its body temperature dips below 0°C (32°F). Results
Periodically, perhaps every two weeks or so, hibernating Day Night
animals undergo arousal, raising their body temperature Per2 Bmal1
and becoming active briefly before resuming hibernation.
100
Relative RNA level (%)

Metabolic rates during hibernation can be 20 times lower
than if the animal attempted to maintain normal body tem- 80
peratures of 36938°C 1979100°F2. As a result, hibernators 60
such as the ground squirrel can survive through the winter 40
on limited supplies of energy stored in the body tissues or as
20
food cached in a burrow. Similarly, the slow metabolism and
inactivity of estivation, or summer torpor, enable animals to 0
Euthermia Hibernation Euthermia Hibernation
survive long periods of high temperatures and scarce water.
What happens to the circadian rhythm in hibernating ani- Conclusion Hibernation disrupted circadian variation
mals? In the past, researchers reported detecting daily biological in the hamster’s clock gene RNA levels. Further experi-
rhythms in hibernating animals. However, in some cases the ments demonstrated that this disruption was not simply
animals were probably in a state of torpor from which they could due to the dark environment during hibernation, since
for nonhibernating animals RNA levels during a darkened
readily arouse, rather than “deep” hibernation. More recently, a daytime were the same as in daylight. The researchers con-
group of researchers in France addressed this question in a differ- cluded that the biological clock stops running in hibernat-
ent way, examining the machinery of the biological clock rather ing European hamsters and, perhaps, in other hibernators
as well.
than the rhythms it controls (Figure 40.22). Working with the
European hamster, they found that molecular components of Data from F. G. Revel et al., The circadian clock stops ticking during deep hiberna-
tion in the European hamster, Proceedings of the National Academy of Sciences
the clock stopped oscillating during hibernation. These findings USA 104:13816–13820 (2007).
support the hypothesis that the circadian clock ceases operation
during hibernation, at least in this species. WHAT IF? Suppose you discovered a new hamster gene and found
that the levels of RNA for this gene were constant during hibernation.
From tissue types to homeostasis, this chapter has focused What could you conclude about the day and night RNA levels for this
on the whole animal. We also investigated how animals gene during euthermia?
exchange materials with the environment and how size and
activity affect metabolic rate. For much of the rest of this unit,
we’ll explore how specialized organs and organ systems
CONCEPT CHECK 40.4
enable animals to meet the basic challenges of life. In Unit 6,
1. If a mouse and a small lizard of the same mass (both at rest)
we investigated how plants meet the same challenges.
were placed in experimental chambers under identical envi-
Figure 40.23, on the next two pages, highlights some funda- ronmental conditions, which animal would consume oxygen
mental similarities and differences in the evolutionary adapta- at a higher rate? Explain.
tions of plants and animals. This figure is thus a review of Unit 2. Which animal must eat a larger proportion of its weight in food
6, an introduction to Unit 7, and, most importantly, an illus- each day: a house cat or an African lion caged in a zoo? Explain.
tration of the connections that unify the myriad forms of life. 3. WHAT IF? Suppose the animals at a zoo were resting com-
fortably and remained at rest while the nighttime air tem-
perature dropped. If the temperature change were sufficient
Mastering Biology Interview with George to cause a change in metabolic rate, what changes would you
Bartholomew: Exploring connections between expect for an alligator and a lion?
animal physiology and the environment
For suggested answers, see Appendix A.

▼ Figure 40.23
MAKE CONNECTIONS
Life Challenges and Solutions
in Plants and Animals
Multicellular organisms face a common set
of challenges. Comparing the solutions that
have evolved in plants and animals reveals
both unity (shared elements) and diversity
(distinct features) across these two lineages.
Nutritional Mode
All living things must obtain energy and carbon from the
environment to grow, survive, and reproduce. Plants are
autotrophs, obtaining their energy through photosynthesis
and their carbon from inorganic sources, whereas animals are
heterotrophs, obtaining their energy and carbon from food.
Evolutionary adaptations in plants and animals support these
different nutritional modes. The broad surface of many leaves
enhances light capture for photosynthesis. When hunting, a
bobcat relies on stealth, speed, and sharp claws. (See Figure 36.2
and Figure 41.16.)
Growth and Regulation

The growth and physiology of both plants and animals are
regulated by hormones. In plants, hormones may act in a local
area or be transported in the body. They control growth patterns,
flowering, fruit development, and more. In animals, hormones
circulate throughout the body and act in specific target tissues,
controlling homeostatic processes and developmental events such
as molting. (See Figure 39.10 and Figure 45.12.)
Environmental Response
All forms of life must detect and respond Mastering Biology BioFlix®
Animation: Homeostasis:
appropriately to conditions in their
Regulating Blood Sugar
environment. Specialized organs sense
environmental signals. For example,
the floral head of a sunflower and an
insect’s eyes both contain photoreceptors
that detect light. Environmental signals
activate specific receptor proteins,
triggering signal transduction pathways
that initiate cellular responses
coordinated by chemical and electrical
communication. (See Figure 39.19 and
Figure 50.15.)

Reproduction
In sexual reproduction,
specialized tissues and
structures produce and
exchange gametes. Offspring
are generally supplied with
nutritional stores that facilitate
Transport rapid growth and development.
All but the simplest For example, seeds have stored
multicellular organisms food reserves that supply energy to the young seedling, while
must transport nutrients and milk provides sustenance for juvenile mammals. (See Figure
waste products between locations in the body. A system of 38.8 and Figure 34.40.)
tubelike vessels is the common evolutionary solution, while the
mechanism of circulation varies. Plants harness solar energy
to transport water, minerals, and sugars through specialized
tubes (left). In animals, a pump (heart) moves circulatory fluid
through vessels (right). (See Figure 35.10 and Figure 42.9.)
Mastering Biology BioFlix® Animation:

Water Transport in Plants
Gas Exchange
The exchange of certain
gases with the environment
is essential for life.
Respiration by plants and
animals requires taking up
oxygen (O2) and releasing carbon dioxide (CO2). In photosynthesis,
net exchange occurs in the opposite direction: CO2 uptake and O2
release. In both plants and animals, highly convoluted surfaces
that increase the area available for gas exchange have evolved,
such as the spongy mesophyll of leaves (left) and the alveoli of
lungs (right). (See Figure 35.18 and Figure 42.24.)
Mastering Biology BioFlix® Animation:

Absorption Gas Exchange
Organisms need to absorb nutrients. The root hairs

of plants (left) and the villi (projections) that line
MAKE CONNECTIONS
the intestines of vertebrates (right) increase the
Compare the adaptations that enable plants and animals to
surface area available for absorption. (See Figure 35.3
respond to the challenges of living in hot and cold environments.
and Figure 41.12.) See Concept 39.4 and Concept 40.3.

40 Chapter Review Go to Mastering Biology for Assignments, the eText,
the Study Area, and Dynamic Study Modules.
• Regulated change in the internal environment is essential to

SUMMARY OF KEY CONCEPTS normal function. Circadian rhythms are daily fluctuations in
metabolism and behavior tuned to the cycles of light and dark
To review key terms, go to the Vocabulary Self-Quiz in the in the environment. Other environmental changes may trigger
Mastering Biology eText or Study Area, or go to goo.gl/zkjz9t. acclimatization, a temporary shift in the steady state.
? Is it accurate to define homeostasis as a constant internal

CONCEPT 40.1 environment? Explain.
Animal form and function are correlated at all levels
of organization (pp. 874–881) CONCEPT 40.3
• Physical laws constrain the evolution of an animal’s size and Homeostatic processes for thermoregulation involve
shape. These constraints contribute to convergent evolution in form, function, and behavior (pp. 884–889)
animal body forms.
• Each animal cell must have access to an aqueous environment. • An animal maintains its internal temperature within a toler-
Simple two-layered sacs and flat shapes maximize exposure to able range by thermoregulation. Endothermic animals are
the surrounding medium. More complex body plans have highly warmed mostly by heat generated by metabolism. Ectothermic
folded internal surfaces specialized for exchanging materials. animals get most of their heat from external sources. Endothermy
• Animal bodies are based on a hierarchy of cells, tissues, organs, requires a greater expenditure of energy. Body temperature may
and organ systems. Epithelial tissue forms active interfaces on vary with environmental temperature, as in poikilotherms, or be
external and internal surfaces; connective tissue binds and sup- relatively constant, as in homeotherms.
ports other tissues; muscle tissue contracts, moving body parts; • In thermoregulation, physiological and behavioral adjust-
and nervous tissue transmits nerve impulses throughout the body. ments balance heat gain and loss, which occur through radia-
• The endocrine and nervous systems are the two means of tion, evaporation, convection, and conduction. Insulation and
communication between different locations in the body. The en- countercurrent exchange reduce heat loss, whereas panting,
docrine system broadcasts signaling molecules called hormones sweating, and bathing increase evaporation, cooling the body.
everywhere via the bloodstream, but only certain cells are respon- Many ectotherms and endotherms adjust their rate of heat
sive to each hormone. The nervous system uses dedicated cellular exchange with their surroundings by vasodilation or vasocon-
circuits involving electrical and chemical signals to send informa- striction and by behavioral responses.
tion to specific locations. • Many mammals and birds adjust their amount of body insula-
tion in response to changes in environmental temperature.
? For a large animal, what challenges would a spherical shape pose for Ectotherms undergo a variety of changes at the cellular level to
carrying out exchange with the environment? acclimatize to shifts in temperature.
• The hypothalamus acts as the thermostat in mammalian regu-
CONCEPT 40.2 lation of body temperature. Fever reflects a resetting of this ther-
mostat to a higher normal range in response to infection.
Feedback control maintains the internal environment
in many animals (pp. 881–883) ? Given that humans thermoregulate, explain why your skin is cooler
than your body core.
• An animal is a regulator if it controls an internal variable and a
conformer if it allows an internal variable to vary with external CONCEPT 40.4
changes. Homeostasis is the maintenance of a steady state de-
spite internal and external changes. Energy requirements are related to animal size,
• Homeostatic mechanisms are usually based on negative activity, and environment (pp. 889–895)
feedback, in which the response reduces the stimulus. In
contrast, positive feedback involves amplification of a stimulus • Animals obtain chemical energy from food, storing it for short-
by the response and often brings about a change in state, such as term use in ATP. The total amount of energy used in a unit of time
the transition from pregnancy to childbirth. defines an animal’s metabolic rate.
• Under similar conditions and for animals of the same size,
the basal metabolic rate of endotherms is substantially
NORMAL RANGE
higher than the standard metabolic rate of ectotherms.
for internal variable
Minimum metabolic rate per gram is inversely related to body
size among similar animals. Animals allocate energy for basal
(or standard) metabolism, activity, homeostasis, growth,
Stimulus: change in and reproduction.
Response • Torpor, a state of decreased activity and metabolism,
internal variable
conserves energy during environmental extremes. Animals
may enter torpor according to a circadian rhythm (daily torpor),
in winter (hibernation), or in summer (estivation).
Control center Sensor
? Why do small animals breathe more rapidly than large animals?

7. Which of the following animals uses the largest percentage of
TEST YOUR UNDERSTANDING its energy budget for homeostatic regulation?
(A) marine jelly (an invertebrate)
For more multiple-choice questions, go to the Practice Test in the (B) snake in a temperate forest
Mastering Biology eText or Study Area, or go to goo.gl/GruWRg. (C) desert insect
(D) desert bird
Levels 1-2: Remembering/Understanding
8. DRAW IT Draw a model of the control circuit(s) required for
1. The body tissue that consists largely of material located outside driving an automobile at a fairly constant speed over a hilly road.
of cells is Indicate each feature that represents a sensor, stimulus, or response.
(A) epithelial tissue.
(B) connective tissue. Levels 5-6: Evaluating/Creating
(C) muscle tissue. 9. EVOLUTION CONNECTION In 1847, the German biologist
(D) nervous tissue. Christian Bergmann noted that mammals and birds living
2. Which of the following would increase the rate of heat at higher latitudes (farther from the equator) are on average
exchange between an animal and its environment? larger and bulkier than related species found at lower latitudes.
(A) feathers or fur Suggest an evolutionary hypothesis to explain this observation.
(B) vasoconstriction 10. SCIENTIFIC INQUIRY Eastern tent caterpillars (Malacosoma
(C) wind blowing across the body surface americanum) live in large groups in silk nests resembling tents,
(D) countercurrent heat exchanger which they build in trees. They are among the first insects to
be active in early spring, when daily temperature fluctuates
3. Consider the energy budgets for a human, an elephant, a from freezing to very hot. Over the course of a day, they display
penguin, a mouse, and a snake. The ________ would have the striking differences in behavior: Early in the morning, they rest
highest total annual energy expenditure, and the ________ in a tightly packed group on the tent’s east-facing surface. In
would have the highest energy expenditure per unit mass. midafternoon, they are on its undersurface, each caterpillar
(A) elephant; mouse hanging by a few of its legs. Propose a hypothesis to explain
(B) elephant; human this behavior. How could you test it?
(C) mouse; snake
11. SCIENCE, TECHNOLOGY, AND SOCIETY Medical
(D) penguin; mouse
researchers are investigating artificial substitutes for various
human tissues. Why might artificial blood or skin be useful?
Levels 3-4: Applying/Analyzing What characteristics would these substitutes need in order to
4. Compared with a smaller cell, a larger cell of the same function well in the body? Why do real tissues work better?
shape has Why not use the real tissues if they work better? What other
(A) less surface area. artificial tissues might be useful? What problems do you
(B) less surface area per unit of volume. anticipate in developing and applying them?
(C) the same surface-area-to-volume ratio. 12. WRITE ABOUT A THEME: ENERGY AND MATTER In a
(D) a smaller cytoplasm-to-nucleus ratio. short essay (about 100–150 words) focusing on energy transfer
and transformation, discuss the advantages and disadvantages
5. An animal’s inputs of energy and materials would exceed
of hibernation.
its outputs
(A) if the animal is an endotherm, which must always take in 13. SYNTHESIZE YOUR KNOWLEDGE
more energy because of its high metabolic rate.
(B) if it is actively foraging for food.
(C) if it is growing and increasing its mass.
(D) never; due to homeostasis, these energy and material bud-
gets always balance.
6. You are studying a large tropical reptile that has a high and
relatively stable body temperature. How do you determine
whether this animal is an endotherm or an ectotherm?
(A) You know from its high and stable body temperature that it
must be an endotherm.
(B) You subject this reptile to various temperatures in the lab
and find that its body temperature and metabolic rate
change with the ambient temperature. You conclude that it
is an ectotherm.
(C) You note that its environment has a high and stable temper-
ature. Because its body temperature matches the environ- These macaques (Macaca fuscata) are partially immersed in a
mental temperature, you conclude that it is an ectotherm. hot spring in a snowy region of Japan. What are some ways that
(D) You measure the metabolic rate of the reptile, and because it form, function, and behavior contribute to homeostasis for
is higher than that of a related species that lives in temper- these animals?
ate forests, you conclude that this reptile is an endotherm
and its relative is an ectotherm. For selected answers, see Appendix A.

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Unit 7 ANIMAL FORM AND

40.1 Animal form and function

40.2 Feedback control maintains the

40.3 Homeostatic processes for

40.4 Energy requirements are related

874 UNIT SEVEN Animal Form and Function

c Figure 40.4 Internal exchange surfaces of EXTERNAL ENVIRONMENT

excretory systems all have such exchange

CHAPTER 40 Basic Principles of Animal Form and Function 875

Table 40.1 Organ Systems in Mammals

Organ System Main Components Main Functions

876 UNIT SEVEN Animal Form and Function

Occurring as sheets of cells, epithelial tissues, or epithelia (singu- Stratified squamous

Cuboidal epithelium Simple columnar Simple squamous Pseudostratified

The single layer of platelike

Lumen Apical surface

CHAPTER 40 Basic Principles of Animal Form and Function 877

from the loose weave of its Plasma

bones at joints. loose connective tissue that fibers embedded in a rubbery

canal containing blood vessels and nerves.

Central canal Chondrocytes

878 UNIT SEVEN Animal Form and Function

Mastering Biology Animation: Muscle Tissue

100 om Nucleus Muscle fibers 25 om Nucleus Intercalated disk 25 om

Mastering Biology Animation: Nervous Tissue

muscles, or other cells

CHAPTER 40 Basic Principles of Animal Form and Function 879

Different hormones cause distinct effects, and only

880 UNIT SEVEN Animal Form and Function

in the face of external fluctuation. In 30

CHAPTER 40 Basic Principles of Animal Form and Function 881

882 UNIT SEVEN Animal Form and Function

(a) Variation in core body temperature and melatonin concentra-

clock is normally coordinated with the cycle of light and

CHAPTER 40 Basic Principles of Animal Form and Function 883

884 UNIT SEVEN Animal Form and Function

CHAPTER 40 Basic Principles of Animal Form and Function 885

tates heat transfer from arteries to

886 UNIT SEVEN Animal Form and Function

Adjusting Metabolic Heat Production

CHAPTER 40 Basic Principles of Animal Form and Function 887

888 UNIT SEVEN Animal Form and Function

NORMAL BODY TEMPERATURE

Body temperature decreases

Blood vessels in skin Thermostat in

CONCEPT CHECK 40.3 reproduction. The overall flow and transformation of

CHAPTER 40 Basic Principles of Animal Form and Function 889

. Figure 40.19 Measuring the rate of oxygen consumption by a swimming shark.

890 UNIT SEVEN Animal Form and Function

BMR (L O2/hr) (log scale)

BMR per kg body mass (L O2/hr • kg)

CHAPTER 40 Basic Principles of Animal Form and Function 891

892 UNIT SEVEN Animal Form and Function

Relative RNA level (%)

CHAPTER 40 Basic Principles of Animal Form and Function 893

Growth and Regulation

894 UNIT SEVEN Animal Form and Function

Mastering Biology BioFlix® Animation:

Mastering Biology BioFlix® Animation: