CONCEPT 34.1 the heart.

The heart powers circulation by using metabolic en-

ergy to elevate the circulatory fluid’s hydrostatic pressure, the
Circulatory systems link exchange pressure the fluid exerts on surrounding vessels. The fluid then
surfaces with cells throughout flows through the vessels and back to the heart.
Circulatory systems are either open or closed. In an open
the body circulatory system, the circulatory fluid, called hemolymph,
The molecular trade that an animal carries out with its is also the interstitial fluid that bathes body cells. Arthropods,
environment involves gaining O2 and nutrients while shed- such as grasshoppers, and some molluscs, such as clams, have
ding CO2 and other waste products. How does this exchange an open circulatory system. Contraction of the heart pumps
take place? Small molecules, including O2 and CO2, undergo the hemolymph through the circulatory vessels into intercon-
random thermal motion—diffusion (see Concept 5.3). When nected sinuses, spaces surrounding the organs (Figure 34.3a).
there is a difference in concentration, such as between a cell There, chemical exchange occurs between the hemolymph
and its immediate surroundings, diffusion can result in net and body cells. Relaxation of the heart draws hemolymph back
movement. But such movement is very slow for distances of in through pores, which have valves that close when the heart
more than a few millimeters. That’s because the time it takes
for a substance to diffuse from one place to another is propor-
tional to the square of the distance. For example, a quantity of
glucose that takes 1 second to diffuse 100 μm will take 100 sec- ▼ Figure 34.3 Open and closed circulatory systems.
onds to diffuse 1 mm and almost 3 hours to diffuse 1 cm. (a) An open circulatory system
Given that net movement by diffusion is rapid only over very
small distances, how does each cell of an animal participate in
Heart
exchange? Natural selection has resulted in two basic adapta-
tions that permit effective exchange for all of an animal’s cells.
One adaptation for efficient exchange is a body that is only
one or two cells thick. Each cell can thus exchange gases di- Hemolymph in sinuses
rectly with the surrounding medium. Such an arrangement is surrounding organs
characteristic of certain invertebrates, including cnidarians,
such as hydras and jellies, and flatworms (Figure 34.2). In
these animals, a central gastrovascular cavity functions in di- Pores
gestion and in distributing throughout the body the nutrients
released from food by digestion.
Animals that lack a simple body plan display an alternative
adaptation for efficient exchange: a circulatory system, which
moves fluid between each cell’s immediate surroundings and the
body tissues. As a result, exchange with the environment and ex- Tubular heart
change with body tissues both occur over very short distances.
(b) A closed circulatory system
Open and Closed Circulatory Systems
A circulatory system has three basic components: a circulatory Heart
fluid, a set of interconnecting vessels, and a muscular pump,
Interstitial fluid
Mouth Gastrovascular cavity Blood
Small branch
vessels in
each organ

Dorsal vessel Auxiliary

1 mm
(main heart) hearts
▲ Figure 34.2 The flat body plan of the planarian Dugesia
(LM). The planarian’s flat shape enables gas exchange to occur across
the entire body surface. A gastrovascular cavity (here stained red) that
branches throughout the body extracts and distributes nutrients from Ventral vessels
food that enters the mouth.

CHAPTER 34 CIRCULATION AND GAS EXCHANGE 707

contracts. Body movements periodically squeeze the sinuses, blood away from the heart toward capillaries, and veins return
helping circulate the hemolymph. blood toward the heart from capillaries.
In a closed circulatory system, a circulatory fluid called The hearts of all vertebrates contain two or more muscular
blood is confined to vessels and is distinct from the interstitial chambers. The chambers that receive blood entering the heart
fluid (Figure 34.3b). This type of circulatory system is found are called atria (singular, atrium). The chambers responsible
in annelids (including earthworms), many molluscs, and all for pumping blood out of the heart are called ventricles. The
vertebrates. In closed circulatory systems, one or more hearts number of chambers and the extent to which they are sepa-
pump blood into large vessels that branch into smaller ones rated from one another differ among groups of vertebrates, as
that infiltrate the tissues and organs. Chemical exchange oc- we’ll discuss next. These important differences reflect the close
curs between the blood and the interstitial fluid, as well as fit of form to function that arises from natural selection.
between the interstitial fluid and body cells.
The fact that both open and closed circulatory systems are Single Circulation
widespread among animals suggests that each system offers In sharks, rays, and bony fishes, blood travels through the body
evolutionary advantages. The lower hydrostatic pressures typi- and returns to its starting point in a single circuit (loop), an
cally associated with open circulatory systems make them less arrangement called single circulation (Figure 34.4a). These
costly than closed systems in terms of energy expenditure. In animals have a heart that consists of two chambers: an atrium
some invertebrates, open circulatory systems serve additional and a ventricle. Blood entering the heart collects in the atrium
functions. For example, spiders use the hydrostatic pressure of before transfer to the ventricle. Contraction of the ventricle
their open circulatory system to extend their legs. pumps blood to a capillary bed in the gills, where there is a net
The benefits of closed circulatory systems include blood diffusion of O2 into the blood and of CO2 out of the blood. As
pressure high enough to enable the effective delivery of O2 blood leaves the gills, the capillaries converge into a vessel that
and nutrients to the cells of larger and more active animals. carries oxygen-rich blood to capillary beds throughout the
Among the molluscs, for instance, closed circulatory systems body. Blood then returns to the heart.
are found in the largest and most active species, the squids In single circulation, blood that leaves the heart passes
and octopuses. Closed systems are also particularly well suited through two capillary beds (in the gills and elsewhere in the
to regulating the distribution of blood to different organs, as body) before returning to the heart. When blood flows through
you’ll learn later in this chapter. In examining closed circula- a capillary bed, blood pressure drops substantially, for reasons
tory systems in more detail, we’ll focus in this section on we’ll explain shortly. The drop in blood pressure in the gills lim-
the vertebrates. its the rate of blood flow in the rest of the animal’s body. As the
animal swims, however, the contraction and relaxation of its
Organization of Vertebrate muscles help accelerate the pace of circulation.
Circulatory Systems
The term cardiovascular system is often used to refer to the Double Circulation
heart and blood vessels of humans and other vertebrates. Blood The circulatory systems of amphibians, reptiles, and mammals
circulates to and from the heart through an amazingly extensive have two circuits of blood flow, an arrangement called double
network of vessels: The total length of blood vessels in an aver- circulation (Figure 34.4b and c). In animals with double circu-
age human adult is twice Earth’s circumference at the equator! lation, the pumps for the two circuits are combined into a single
Arteries, veins, and capillaries are the three main types of organ, the heart. Having both pumps within a single heart
blood vessels. Within each type, blood flows in one direction. simplifies coordination of the pumping cycles. One pump,
Arteries carry blood from the heart to organs throughout the the right side of the heart, delivers oxygen-poor blood to the
body. Within organs, arteries branch into arterioles. These capillary beds of the gas exchange tissues, where there is a net
small vessels convey blood to capillaries, microscopic ves- movement of O2 into the blood and of CO2 out of the blood.
sels with very thin, porous walls. Local networks of capillaries, This part of the circulation is called a pulmocutaneous circuit if
called capillary beds, infiltrate tissues, passing within a few cell it includes capillaries in both the lungs and the skin, as in many
diameters of every cell in the body. Across the thin walls of amphibians. It is called a pulmonary circuit if the capillary beds
capillaries, chemicals, including dissolved gases, are exchanged involved are all in the lungs, as in reptiles and mammals.
by diffusion between the blood and the interstitial fluid around After the oxygen-enriched blood leaves the gas exchange
the tissue cells. At their “downstream” end, capillaries converge tissues, it enters the other pump, the left side of the heart.
into venules, and venules converge into veins, the vessels that Heart contraction propels this blood to capillary beds in or-
carry blood back to the heart. gans and tissues throughout the body. Following the exchange
Note that arteries and veins are distinguished by the of O2 and CO2, as well as nutrients and waste products, the
direction in which they carry blood, not by the O2 content or now oxygen-poor blood returns to the heart, completing the
other characteristics of the blood they contain. Arteries carry systemic circuit.

708 UNIT SIX ANIMAL FORM AND FUNCTION

 ▼ Figure 34.4 Examples of vertebrate circulatory system organization.

(a) Single circulation: fish (b) Double circulation: amphibian (c) Double circulation: mammal

Gill Pulmocutaneous circuit Pulmonary circuit

capillaries
Lung Lung
and skin capillaries
capillaries
Artery
Atrium Atrium
(A) (A)
Heart:
A A
Atrium (A)
Ventricle (V) V V
Right Left Right Left
Vein Ventricle (V)

Systemic Systemic
capillaries capillaries
Body
capillaries Systemic circuit Systemic circuit

Key Oxygen-rich blood (Note that circulatory systems are shown as if the body were facing you: The right side
Oxygen-poor blood of the heart is shown on the left, and vice versa.)

Double circulation provides a vigorous flow of blood to the t
In the three-chambered heart of turtles, snakes, and lizards,
brain, muscles, and other organs because the heart repressur- an incomplete septum partially divides the single ventricle
izes the blood destined for these tissues after it passes through into right and left chambers. Two major arteries, called aor-
the capillary beds of the lungs or skin. Indeed, blood pressure tas, lead to the systemic circulation. As with amphibians, the
is often much higher in the systemic circuit than in the gas circulatory system enables control of the relative amount of
exchange circuit. By contrast, in single circulation the blood blood flowing to the lungs and the rest of the body.
flows under reduced pressure directly from the gas exchange t
In alligators, caimans, and other crocodilians, the ventricles
organs to other organs. are divided by a complete septum, but the pulmonary and
systemic circuits connect where the arteries exit the heart.
Evolutionary Variation in Double Circulation This connection allows arterial valves to shunt blood flow
EVOLUTION Some vertebrates with double circulation are in- away from the lungs temporarily, such as when the animal
termittent breathers. For example, amphibians and many rep- is underwater.
tiles fill their lungs with air periodically, passing long periods
Double circulation in birds and mammals, which for the
either without gas exchange or by relying on another gas ex-
most part breathe continuously, differs from double circula-
change tissue, typically the skin. A variety of adaptations found
tion in other vertebrates. As shown for a panda in Figure 34.4c,
among intermittent breathers enable their circulatory systems
the heart has two atria and two completely divided ventricles.
to temporarily bypass the lungs in part or in whole:
The left side of the heart receives and pumps only oxygen-rich
t
Frogs and other amphibians have a heart with three blood, while the right side receives and pumps only oxygen-
chambers—two atria and one ventricle (see Figure 34.4b). poor blood. Unlike amphibians and many reptiles, birds and
A ridge within the ventricle diverts most (about 90%) of the mammals cannot vary blood flow to the lungs without varying
oxygen-rich blood from the left atrium into the systemic cir- blood flow throughout the body in parallel.
cuit and most of the oxygen-poor blood from the right atrium How has natural selection shaped the double circulation of
into the pulmocutaneous circuit. When a frog is underwater, birds and mammals? As endotherms, they use about ten times
the incomplete division of the ventricle allows the frog to adjust as much energy as equal-sized ectotherms (see Concept 33.5).
its circulation, largely shutting off blood flow to its temporarily Their circulatory systems therefore need to deliver about ten
ineffective lungs. Blood flow continues to the skin, which acts times as much fuel and O2 to their tissues and remove ten
as the sole site of gas exchange while the frog is submerged. times as much CO2 and other wastes. This large traffic of

CHAPTER 34 CIRCULATION AND GAS EXCHANGE 709

substances is made possible by the separate and independently Superior Capillaries of
vena cava 7 head and
powered systemic and pulmonary circuits and by large hearts.
forelimbs
A powerful four-chambered heart arose independently in the
distinct ancestors of birds and mammals and thus reflects con- Pulmonary Pulmonary
vergent evolution. artery artery
In the next section, we’ll restrict our focus to circulation in
mammals and to the anatomy and physiology of the key circu- Capillaries 9 Aorta Capillaries
latory organ—the heart. of right lung of left lung
6

CONCEPT CHECK 34.1

1. How is the flow of hemolymph through an open circula- 2 3
3
tory system similar to the flow of water through an outdoor 4
11
fountain?
2. Three-chambered hearts with incomplete septa were once Pulmonary Pulmonary
viewed as being less adapted to circulatory function than 5
vein 1 vein
mammalian hearts. What advantage of such hearts did this Right atrium 10 Left atrium
viewpoint overlook?
Right ventricle Left ventricle
3. WHAT IF? The heart of a normally developing human fetus
has a hole between the left and right atria. In some cases, Inferior Aorta
this hole does not close completely before birth. If the hole vena cava
weren’t surgically corrected, how would it affect the O2 con-
tent of the blood entering the systemic circuit? Capillaries of
For suggested answers, see Appendix A. abdominal organs
8 and hind limbs

▲ Figure 34.5 The mammalian cardiovascular system: an

CONCEPT 34.2 overview. Note that the dual circuits operate simultaneously, not in
the serial fashion that the numbering in the diagram suggests. The
Coordinated cycles of heart two ventricles contract almost in unison and pump the same volume of
blood. However, the systemic circuit has a much greater total volume
contraction drive double circulation than the pulmonary circuit.
in mammals
The timely delivery of O2 to the body’s organs is critical: Some arteries (not shown), which supply blood to the heart muscle
brain cells, for example, die if their O2 supply is interrupted itself. Then branches lead to 7 capillary beds in the head and
for even a few minutes. How does the mammalian cardiovas- arms (forelimbs). The aorta then descends into the abdomen,
cular system meet the body’s continuous (although variable) supplying oxygen-rich blood to arteries leading to 8 capillary
demand for O2? To answer this question, we must consider beds in the abdominal organs and legs (hind limbs). Within the
how the parts of the system are arranged and how each capillary beds, there is a net diffusion of O2 from the blood to
part functions. the tissues and of CO2 (produced by cellular respiration) into
the blood. Capillaries rejoin, forming venules, which convey
Mammalian Circulation blood to veins.
Let’s first examine the overall organization of the mammalian As it moves back toward the heart, oxygen-poor blood from
cardiovascular system, beginning with the pulmonary cir- the head, neck, and forelimbs is channeled into a large vein,
cuit. (The circled numbers refer to corresponding locations 9 the superior vena cava. Another large vein, 10 the inferior
in Figure 34.5.) Contraction of 1 the right ventricle pumps vena cava, drains blood from the trunk and hind limbs. The two
blood to the lungs via 2 the pulmonary arteries. As the blood venae cavae empty their blood into 11 the right atrium, from
flows through 3 capillary beds in the left and right lungs, it which the oxygen-poor blood flows into the right ventricle.
loads O2 and unloads CO2. Oxygen-rich blood returns from
the lungs via the pulmonary veins to 4 the left atrium of the The Mammalian Heart: A Closer Look
heart. Next, the oxygen-rich blood flows into 5 the heart’s left Located behind the sternum (breastbone), the human heart
ventricle, which pumps the oxygen-rich blood out to body tis- is about the size of a clenched fist and consists mostly of car-
sues through the systemic circuit. diac muscle. The two atria have relatively thin walls and serve
Blood leaves the left ventricle of the heart via 6 the aorta, as collection chambers for blood returning to the heart from
which conveys blood to arteries leading throughout the body. the lungs or other body tissues (Figure 34.6). Much of the
The first branches leading from the aorta are the coronary blood that enters the atria flows into the ventricles while all

710 UNIT SIX ANIMAL FORM AND FUNCTION

Pulmonary artery Aorta
2 Atrial systole and ventricular
Pulmonary diastole. A brief period of atrial
artery contraction then forces all blood
remaining in the atria into the
Right ventricles.
atrium 1 Atrial and
Left ventricular diastole.
atrium During a relaxation
phase, blood returning
from the large veins
flows into the atria and
then into the ventricles
through the AV valves.
Semilunar Semilunar
valve valve

Atrioventricular Atrioventricular
(AV) valve (AV) valve
0.1 sec
Right Left
ventricle ventricle
0.3 sec
▲ Figure 34.6 The mammalian heart: a closer look. Notice 0.4 sec
the locations of the valves, which prevent backflow of blood within the
heart. Also notice how the atria and left and right ventricles differ in
the thickness of their muscular walls.

heart chambers are relaxed. The remainder is transferred by

contraction of the atria before the ventricles begin to contract. 3 Ventricular systole and atrial
Compared to the atria, the ventricles have thicker walls and diastole. During the remainder of
the cycle, ventricular contraction
contract much more forcefully—especially the left ventricle, pumps blood into the large
which pumps blood throughout the body via the systemic arteries through the semilunar
circuit. Although the left ventricle contracts with greater force valves.
than the right ventricle, it pumps the same volume of blood as
▲ Figure 34.7 The cardiac cycle. Note that during all but
the right ventricle during each contraction.
0.1 second of the cardiac cycle, the atria are relaxed and are filling
The heart contracts and relaxes in a rhythmic cycle. When with blood returning via the veins.
it contracts, it pumps blood; when it relaxes, its chambers fill
with blood. One complete sequence of pumping and filling is
referred to as the cardiac cycle (Figure 34.7). The contraction the ventricles closes the AV valves, keeping blood from flowing
phase of the cycle is called systole (pronounced sis9-tō-lē), and back into the atria. Semilunar valves are located at the two
the relaxation phase is called diastole (dī-as9-tō-lē). exits of the heart: where the pulmonary artery leaves the right
The volume of blood each ventricle pumps per minute is ventricle and where the aorta leaves the left ventricle. These
the cardiac output. Two factors determine cardiac output: valves are pushed open by the pressure generated during con-
the rate of contraction, or heart rate (beats per minute), traction of the ventricles. When the ventricles relax, blood
and the stroke volume, the amount of blood pumped by a pressure built up in the pulmonary artery and aorta closes the
ventricle in a single contraction. The average stroke volume semilunar valves and prevents significant backflow.
in humans is about 70 mL. Multiplying this stroke volume You can follow the closing of the two sets of heart valves
by a resting heart rate of 72 beats per minute yields a car- either with a stethoscope or by pressing your ear tightly against
diac output of 5 L/min—about equal to the total volume of the chest of a friend (or a friendly dog). The sound pattern is
blood in the human body. During heavy exercise, when O2 “lub-dup, lub-dup, lub-dup.” The first heart sound (“lub”) is
demand is especially high, cardiac output increases as much created by the recoil of blood against the closed AV valves. The
as fivefold. second sound (“dup”) is due to the vibrations caused by closing
Four valves in the heart prevent backflow and keep blood of the semilunar valves.
moving in the correct direction (see Figures 34.6 and 34.7). If blood squirts backward through a defective valve, it may
Made of flaps of connective tissue, the valves open when produce an abnormal sound called a heart murmur. Some
pushed from one side and close when pushed from the other. people are born with heart murmurs; in others, the valves may
An atrioventricular (AV) valve lies between each atrium and be damaged by infection (for instance, from rheumatic fever,
ventricle. Pressure generated by the powerful contraction of an inflammation of the heart or other tissues triggered by

CHAPTER 34 CIRCULATION AND GAS EXCHANGE 711

1 Signals (yellow) 2 Signals are delayed at 3 Bundle branches 4 Signals spread the left and right atria. Here the impulses
from SA node spread AV node. pass signals to throughout are delayed for about 0.1 second before
through atria. heart apex. ventricles.
spreading to the heart apex. This delay
allows the atria to empty completely
before the ventricles contract. Then the
signals from the AV node are conducted
to the heart apex and throughout the
ventricular walls.
Physiological cues alter heart tempo
SA node AV
(pacemaker) node by regulating the pacemaker function
Bundle Purkinje of the SA node. For example, when you
branches Heart fibers
apex stand up and start walking, the nervous
system speeds up your pacemaker. The
ECG resulting increase in heart rate provides
the additional O2 needed by the muscles
▲ Figure 34.8 The control of heart rhythm. Electrical signals follow a set path through the that are powering your activity. If you
heart in establishing the heart rhythm. The diagrams at the top trace the movement of electrical then sit down and relax, the nervous
signals (yellow) during the cardiac cycle; specialized cells involved in electrical control of the rhythm system slows down your pacemaker,
are indicated in orange. Under each step, the corresponding portion of an electrocardiogram (ECG)
is highlighted in yellow. In step 4, the portion of the ECG to the right of the “spike” represents decreasing your heart rate and thus
electrical activity that reprimes the ventricles for the next round of contraction. conserving energy. Hormones and tem-
WHAT IF? If your doctor gave you a copy of your ECG recording over many cardiac cycles, perature also influence the pacemaker.
how could you determine what your heart rate had been during the test? For instance, epinephrine, the “fight-or-
flight” hormone secreted by the adrenal
infection with certain bacteria). When a valve defect is severe glands, causes the heart rate to increase, as does an increase in
enough to endanger health, surgeons may implant a mechani- body temperature.
cal replacement valve. However, not all heart murmurs are Having examined the operation of the circulatory pump, we
caused by a defect, and most valve defects do not reduce the turn in the next section to the forces and structures that influ-
efficiency of blood flow enough to warrant surgery. ence blood flow in the vessels of each circuit.

Maintaining the Heart’s Rhythmic Beat CONCEPT CHECK 34.2

1. Explain why blood in the pulmonary veins has a higher
In vertebrates, the heartbeat originates in the heart itself. O2 concentration than in the venae cavae, which are
Some cardiac muscle cells are autorhythmic, meaning also veins.
they can contract and relax repeatedly without any signal 2. Why is it important that the AV node delay the electrical
from the nervous system. A group of such cells forms the impulse moving from the SA node and the atria to the
sinoatrial (SA) node, or pacemaker, which sets the rate ventricles?
and timing at which all cardiac muscle cells contract. (In 3. WHAT IF? Suppose that after you exercise regularly for
contrast, some arthropods have pacemakers located in the several months, your resting heart rate decreases, but your
nervous system.) cardiac output at rest is unchanged. What other change in
The SA node produces electrical impulses much like those the function of your heart at rest must have occurred?
produced by nerve cells. Because cardiac muscle cells are elec- For suggested answers, see Appendix A.

trically coupled through gap junctions (see Figure 4.27), im-

pulses from the SA node spread rapidly within heart tissue. In
addition, these impulses generate currents that are conducted CONCEPT 34.3
to the skin via body fluids. In an electrocardiogram (ECG or,
often, EKG, from the German spelling), these currents are re- Patterns of blood pressure and
corded by electrodes placed on the skin. The resulting graph of flow reflect the structure and
current against time has a characteristic shape that represents arrangement of blood vessels
the stages in the cardiac cycle (Figure 34.8).
Impulses from the SA node first spread rapidly through the The vertebrate circulatory system enables blood to deliver
walls of the atria, causing both atria to contract in unison. Dur- oxygen and nutrients and remove wastes throughout the body.
ing atrial contraction, the impulses reach a relay point called In doing so, the circulatory system relies on blood vessels that
the atrioventricular (AV) node, located in the wall between exhibit a close match of structure and function.

712 UNIT SIX ANIMAL FORM AND FUNCTION