Blood Vessel Chapter - NEW

19 The Cardiovascular System:
Blood Vessels
In this chapter, you will learn that
Blood vessels are dynamic structures that control the delivery of blood to body tissues
by exploring
Part 1 Blood Vessel Part 2 Physiology Part 3 Circulatory

Structure and Function of Circulation Pathways
by examining by asking by exploring
19.1 Structure of 19.11 Principal vessels of

blood vessel walls the systemic circulation
19.6 How are flow, 19.9 How is blood
forms
pressure, and flow through
resistance related? tissues controlled?
19.2 Arteries and Arteries Veins
looking closer at
19.7 What is blood and finally, exploring

19.3 Capillaries pressure and how 19.10 Capillary
does it differ in exchange
Developmental Aspects
arteries, capillaries, of Blood Vessels
19.4 Veins and veins?
some form
and
19.5 Anastomoses 19.8 How is blood

pressure regulated?
Neural controls
Ch. 14
Short-term Long-term Renal mechanisms
control control Ch. 26
Hormonal controls
Ch. 16
CAREER CONNECTION
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738
Chapter 19 The Cardiovascular System: Blood Vessels 739
Blood vessels are sometimes compared to a system of ●● In the pulmonary circulation, the arteries, still defined as the
pipes with blood circulating in them, but this analogy is only vessels leading away from the heart, carry oxygen-poor blood
a starting point. Unlike rigid pipes, blood vessels are dynamic to the lungs, and the veins carry oxygen-rich blood from the
structures that pulsate, constrict, relax, and even proliferate lungs to the heart.
(multiply). In this chapter we examine the structure and func- ●● In the special umbilical vessels of a fetus, the roles of veins
tion of these important circulatory passageways. and arteries also differ with respect to oxygenation.
The blood vessels of the body form a closed delivery system
Of all the blood vessels, only the capillaries have intimate
that begins and ends at the heart. The idea that blood circulates
contact with tissue cells and directly serve cellular needs.
in the body dates back to the 1620s with the inspired experi-
Exchanges between the blood and tissue cells occur primarily
ments of William Harvey, an English physician. Prior to that
through the gossamer-thin capillary walls.
time, people thought, as proposed by the ancient Greek physi-
Figure 19.1 summarizes how these vascular channels relate
cian Galen, that blood moved through the body like an ocean
to one another and to vessels of the lymphatic system. The lym-
tide, first moving out from the heart and then ebbing back in
phatic system recovers fluids that leak from the blood vessels
the same vessels.
and is described in Chapter 20.
PART 1
Venous system Arterial system
BLOOD VESSEL
STRUCTURE Large veins Heart
AND FUNCTION (capacitance
vessels)
The three major types of blood vessels are Elastic
arteries, capillaries, and veins. As the heart Large arteries
(pressure
contracts, it forces blood into the large lymphatic
reservoirs)
arteries leaving the ventricles. The blood vessels
then moves into successively smaller
arteries, finally reaching their smallest
Lymph
branches, the arterioles (ar-te9re-ōlz; “little node Muscular
arteries”), which feed into the capillary arteries
beds of body organs and tissues. Blood Lymphatic (distributing
system arteries)
drains from the capillaries into venules
(ven9ūlz), the smallest veins, and then on
into larger and larger veins that merge to Small veins
(capacitance
form the large veins that ultimately empty vessels)
into the heart. Altogether, the blood ves- Arteriovenous
sels in the adult human stretch for about anastomosis
100,000 km (60,000 miles) through the
19
17
internal body landscape!
Lymphatic capillaries
Arteries carry blood away from the
heart, so they are said to “branch,” “diverge,”
or “fork” as they form smaller and smaller Arterioles
divisions. Veins, by contrast, carry blood (resistance
toward the heart and so are said to “join,” vessels)
“merge,” and “converge” into the succes- Postcapillary Terminal
sively larger vessels approaching the heart. venule arteriole
In the systemic circulation, arteries always Capillaries (exchange vessels)
carry oxygenated blood and veins always
carry oxygen-poor blood. The opposite is Figure 19.1 The relationship of blood vessels to each other and to lymphatic
true in two special locations: vessels. Lymphatic vessels recover excess tissue fluid and return it to the blood.
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740 UNIT 4 Maintenance of the Body
(a)
Artery Vein
Artery Vein
Tunica intima
• Endothelium
• Subendothelial layer
• Internal elastic membrane
Tunica media
(smooth muscle and
elastic ﬁbers)
• External elastic membrane
Tunica externa
(collagen ﬁbers)
• Vasa vasorum
Valve
Capillary network
Lumen
19
17 Lumen
Basement membrane
Capillary
Endothelial cells
(b)
Figure 19.2 Generalized structure of arteries, veins, and capillaries. Practice Histology questions: >
(a) Light photomicrograph of a muscular artery and the corresponding vein in cross Study Area > Lab Tools > PAL
section (63). (b) Comparison of wall structure of arteries, veins, and capillaries.
Note that the tunica media is thicker than the tunica externa in arteries and that the
opposite is true in veins.
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19.1 Most blood vessel walls have Table 19.1 Summary of Blood Vessel Anatomy
VESSEL TYPE/ AVERAGE RELATIVE TISSUE
three layers ILLUSTRATION* LUMEN MAKEUP
DIAMETER (D)
Learning Outcomes AND WALL
NN Describe the three layers that typically form the wall of a THICKNESS (T)
us)
blood vessel, and state the function of each.
ageno
NN Define vasoconstriction and vasodilation.
s c le
s
s
Tis s u e
Tis s u e
h Mus
h e li u m
s (C o ll
The walls of all blood vessels, except the very smallest, have
three distinct layers, or tunics (“coverings”), that surround a
Smoot
E la s t ic
F ib r o u
Endot
central blood-containing space, the vessel lumen (Figure 19.2).
The innermost tunic is the tunica intima (in9tĭ-mah). The
name is easy to remember once you know that this tunic is in
ARTERIES
intimate contact with the blood in the lumen. The tunica intima
contains the endothelium, the simple squamous epithelium that
lines the lumen of all vessels ( p. 153). The endothelium is
continuous with the endocardial lining of the heart, and its flat
cells fit closely together, forming a slick surface that minimizes
friction as blood moves through the lumen. In vessels larger D: 1.5 cm
than 1 mm in diameter, a subendothelial layer, consisting of a T: 1.0 mm
Elastic artery
basement membrane and loose connective tissue, supports the
endothelium.
The middle tunic, the tunica media (me9de-ah), is mostly cir-
cularly arranged smooth muscle cells and sheets of elastin. The
activity of the smooth muscle is regulated by sympathetic vaso-
D: 6.0 mm
motor nerve fibers of the autonomic nervous system and a whole
T: 1.0 mm
battery of chemicals. Depending on the body’s needs at any given
Muscular artery
moment, regulation causes either vasoconstriction (lumen diam-
eter decreases as the smooth muscle contracts) or vasodilation
(lumen diameter increases as the smooth muscle relaxes). The
activities of the tunica media are critical in regulating circulatory
dynamics because small changes in vessel diameter greatly influ- D: 37.0 μm
ence blood flow and blood pressure. Generally, the tunica media T: 6.0 μm
is the bulkiest layer in arteries, which bear the chief responsibility Arteriole
for maintaining blood pressure and circulation.
CAPILLARIES
The outermost layer of a blood vessel wall, the tunica externa
(also called the tunica adventitia; ad0ven-tish9e-ah; “coming from
outside”), is composed largely of loosely woven collagen fibers D: 9.0 μm 19
17
that protect and reinforce the vessel, and anchor it to surrounding T: 0.5 μm
structures. The tunica externa is infiltrated with nerve fibers, lym- VEINS
phatic vessels, and, in larger veins, a network of elastic fibers. In
larger vessels, the tunica externa contains a system of tiny blood
vessels, the vasa vasorum (va9sah va-sor9um)—literally, “ves-
sels of the vessels”—that nourish the more external tissues of the D: 20.0 μm
T: 1.0 μm
blood vessel wall. The innermost (luminal) portion of the vessel
Venule
obtains nutrients directly from blood in the lumen.
The three vessel types vary in length, diameter, wall thick-
ness, and tissue makeup (see Table 19.1).
Check Your Understanding

1. Which branch of the autonomic nervous system innervates D: 5.0 mm
blood vessels? Which layer of the blood vessel wall do these T: 0.5 mm
nerves innervate? What are the effectors (cells that carry out Vein
the response)?
*Size relationships are not proportional. Smaller vessels are drawn
2. When vascular smooth muscle contracts, what happens to the relatively larger so detail can be seen. See column 2 for actual
diameter of the blood vessel? What is this called? dimensions.
For answers, see Answers Appendix.
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(Table 19.1). For this reason, they are more active in vasocon-
19.2 Arteries are pressure reservoirs, striction and less capable of stretching. In muscular arteries,
distributing vessels, or resistance however, there is an elastic membrane on each face of the
tunica media.
vessels
Learning Outcome Arterioles
NN Compare and contrast the structure and function of the The smallest of the arteries, arterioles have a lumen diameter
three types of arteries. ranging from 0.3 mm down to 10 μm. Larger arterioles have
all three tunics, but their tunica media is chiefly smooth mus-
In terms of relative size and function, arteries can be divided into cle with a few scattered elastic fibers. Smaller arterioles, which
three groups—elastic arteries, muscular arteries, and arterioles. lead into the capillary beds, are little more than a single layer
of smooth muscle cells spiraling around the endothelial lining.
Elastic Arteries Minute-to-minute blood flow into the capillary beds is deter-
Elastic arteries are the thick-walled arteries near the heart— mined by arteriolar diameter, which varies in response to chang-
the aorta and its major branches (Figure 19.1). These arteries ing neural, hormonal, and local chemical influences. Changing
are the largest in diameter, ranging from 2.5 cm to 1 cm, and diameter changes resistance to blood flow, and so arterioles are
the most elastic (Table 19.1). Because their large lumens make called resistance vessels. When arterioles constrict, the tissues
them low-resistance pathways that conduct blood from the heart served are largely bypassed. When arterioles dilate, blood flow
to medium-sized arteries, elastic arteries are sometimes called into the local capillaries increases dramatically.
conducting arteries.
Elastic arteries contain more elastin than any other vessel Check Your Understanding
type. It is present in all three tunics, but the tunica media con- 3. Name the type of artery that matches each description: major
tains the most. There the elastin constructs concentric “holey” role in dampening the pulsatile pressure of heart contractions;
sheets of elastic connective tissue that look like slices of Swiss vasodilation or constriction determines blood flow to individual
cheese sandwiched between layers of smooth muscle cells. capillary beds; have the thickest tunica media relative to their
Although elastic arteries also contain substantial amounts of lumen size.
smooth muscle, they are relatively inactive in vasoconstriction. For answers, see Answers Appendix.
Thus, in terms of function, they can be visualized as simple
elastic tubes.
Elastic arteries are pressure reservoirs, expanding and 19.3 Capillaries are exchange vessels
recoiling as the heart ejects blood. Consequently, blood flows
Learning Outcome
fairly continuously rather than starting and stopping with the
pulsating rhythm of the heartbeat. If the blood vessels become NN Describe the structure and function of a capillary bed.
hard and unyielding, as in atherosclerosis, blood flows more The microscopic capillaries are the smallest blood vessels
intermittently, similar to the way water flows through a hard (Figure 19.2b). Their exceedingly thin walls consist of just a thin
rubber garden hose attached to a faucet. When the faucet is on, tunica intima surrounded by a basement membrane ( p. 150).
the high pressure makes the water gush out of the hose. But In some cases, a single endothelial cell forms the entire circum-
19
17 when the faucet is shut off, the water flow abruptly becomes ference of the capillary wall. At strategic locations along the
a trickle and then stops, because the hose walls cannot recoil outer surface of some capillaries are spider-shaped pericytes,
to keep the water under pressure. Also, without the pressure- contractile stem cells that can generate new vessels or scar tis-
smoothing effect of the elastic arteries, the walls of arteries sue, stabilize the capillary wall, and help control capillary per-
throughout the body experience higher pressures. Battered by meability (Figure 19.3a).
high pressures, the arteries eventually weaken and may bal- Average capillary length is 1 mm and average lumen diam-
loon out (as an aneurysm) or even burst (see A Closer Look eter is 8–10 μm, just large enough for red blood cells to slip
on p. 792). through in single file. Most tissues have a rich capillary sup-
ply, but there are exceptions. Tendons and ligaments are poorly
Muscular Arteries vascularized (and so heal poorly). Cartilage and epithelia lack
Distally the elastic arteries give way to the muscular arteries, capillaries but receive nutrients from blood vessels in nearby
which deliver blood to specific body organs (and so are some- connective tissues, and the avascular cornea and lens of the eye
times called distributing arteries). Muscular arteries account receive nutrients from the aqueous humor.
for most of the named arteries studied in the anatomy labora- If we compare arteries and arterioles to expressways and
tory. Their internal diameter ranges from that of a little finger to roads, capillaries are the back alleys and driveways that pro-
that of a pencil lead. vide direct access to nearly every cell in the body. Given their
Proportionately, muscular arteries have the thickest tunica location and thin walls, capillaries are ideally suited for their
media of all vessels. Their tunica media contains relatively more role—exchange of materials (gases, nutrients, hormones, and
smooth muscle and less elastic tissue than do elastic arteries so on) between the blood and the interstitial fluid. We describe
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(a) Continuous capillary
Continuous capillaries are the least permeable and most common. Pericyte
• Abundant in skin, muscles, lungs, and CNS. Red blood

• Often have associated pericytes. cell in lumen
• Pinocytotic vesicles ferry fluid across the endothelial cell.
• Most continuous capillaries have intercellular clefts between Basement
endothelial cells. However, brain capillary endothelial cells membrane
lack intercellular clefts and have tight junctions around their
entire perimeter. (This is the structural basis of the blood Endothelial
brain barrier described in Chapter 12.) cell
Intercellular
cleft
Tight junction
Pinocytotic
Endothelial nucleus vesicles
(b) Fenestrated capillary
Fenestrated capillaries have large fenestrations (pores) Pinocytotic

that increase permeability. vesicles
Red blood
• Occur in areas of active filtration (e.g., kidney) or absorption cell in lumen
(e.g., small intestine), and areas of endocrine hormone secretion.
• Fenestrations are Swiss cheese–like holes that tunnel through Fenestrations
endothelial cells. (pores)
• Fenestrations are usually covered by a very thin layer of condensed
Intercellular
extracellular glycoproteins. This layer has little effect on solute and
cleft
fluid movement.
• In some digestive tract organs, the number of fenestrations in Endothelial Endothelial
capillaries increases during active absorption of nutrients. nucleus cell
Basement membrane Tight junction
(c) Sinusoid capillary

Sinusoid capillaries are the most permeable and
occur in limited locations. Endothelial
cell
• Occur in liver, bone marrow, spleen, and adrenal medulla.

Red blood
• Have large intercellular clefts as well as fenestrations; cell in lumen
few tight junctions. 19
17
• Have incomplete basement membranes. Large
intercellular
• Are irregularly shaped and have larger lumens than other capillaries.
cleft
• Allow large molecules and even cells to pass across their walls.
• Blood flows slowly through their tortuous channels.
• Macrophages may extend processes through the clefts to catch Tight junction
Nucleus of
“prey” or, in liver, form part of the sinusoid wall. Incomplete endothelial
basement membrane cell
Figure 19.3 Capillary structure.
these exchanges later in this chapter. Here, we focus on capil- Figure 19.3, notice that all three types have tight junctions that
lary structure. join their endothelial cells together. However, these junctions
are usually incomplete and leave gaps of unjoined membrane
Types of Capillaries called intercellular clefts, which allow limited passage of flu-
Structurally, there are three types of capillaries—continuous, ids and small solutes. Leakier capillaries have specialized pas-
fenestrated, and sinusoid. As you study their properties in sageways that increase fluid movement.
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Capillary Beds Vascular shunt

Precapillary
Capillaries do not function independently. Instead they form sphincters Metarteriole Thoroughfare
interweaving networks called capillary beds. The flow of (closed) channel
blood from an arteriole to a venule—that is, through a capillary
bed—is called the microcirculation. In most body regions, a
terminal arteriole branches into 10 to 20 capillaries (exchange
vessels) that form the capillary bed. These then drain into a
postcapillary venule (Figure 19.4). Blood flow through the
capillary bed is controlled by the diameter of the terminal arteriole
as well as by all of the arterioles upstream from it. As blood
flows through the capillaries, it takes part in exchanges of gases,
nutrients, and wastes with the surrounding tissue cells. True
capillaries
Local chemical conditions and arteriolar vasomotor nerve
Precapillary
fibers regulate the amount of blood entering a capillary bed. A Arteriole sphincters (open) Venule
bed may be flooded with blood or almost completely bypassed,
depending on conditions in the body or in that specific organ. Figure 19.5 Anatomy of a special (mesenteric) capillary
For example, suppose you have just eaten and are sitting bed.
relaxed, listening to your favorite musical group. Food is being
digested, and blood is circulating freely through the capillaries
of your gastrointestinal organs to receive the breakdown prod- capillary beds of your skeletal muscles where it is more immedi-
ucts of digestion. Between meals, however, most of these same ately needed. This rerouting helps explain why vigorous exercise
capillary pathways are closed. right after a meal can cause indigestion or abdominal cramps.
To take another example, when you exercise vigorously, blood In the serous membranes that hold the intestines in place
is rerouted from your digestive organs (food or no food) to the (mesenteries), there are two additional features that form a spe-
cial arrangement of capillaries (Figure 19.5):
●● There is a vascular shunt that directly connects the terminal
arteriole to the postcapillary venule, so that blood can bypass
the true capillaries. This shunt consists of a metarteriole and
a thoroughfare channel.
●● As it branches from the metarteriole, each true capillary is
surrounded by a cuff of smooth muscle called a precapillary
sphincter. The precapillary sphincter acts as a valve to regu-
late blood flow into the capillary. The precapillary sphincters
Terminal
are not innervated, and so are controlled only by local chem-
arteriole Postcapillary ical conditions.
venule
19
17 Capillaries
Arteriole Venule Check Your Understanding
(a) Arterioles dilated—blood flows through capillaries. 4. APPLY Look at Figure 19.4 and assume that the capillary bed
depicted is in your calf muscle. Which condition—(a) or (b)—
would the bed be in if you were doing calf raises at the gym?
Veins are blood reservoirs that

19.4
return blood toward the heart
Learning Outcome
NN Describe the structure and function of veins, and explain
how veins differ from arteries.
Veins carry blood from the capillary beds toward the heart.
(b) Arterioles constricted—no blood flows through capillaries. Along the route, the diameter of successive venous vessels
increases, and their walls gradually thicken as they progress
Figure 19.4 Anatomy of a typical capillary bed.
from venules to larger and larger veins.
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Venules is low. However, the low-pressure condition demands several

structural adaptations to ensure that veins return blood to the
Capillaries unite to form venules, which range from 8 to 100 μm
heart at the same rate it was pumped into the circulation. One
in diameter. The smallest venules, the postcapillary venules,
such adaptation is their large-diameter lumens, which offer rela-
consist entirely of endothelium around which pericytes congre-
tively little resistance to blood flow.
gate. Postcapillary venules are extremely porous (more like cap-
illaries than veins in this way), and fluid and white blood cells Venous Valves
move easily from the bloodstream through their walls. Indeed,
Venous valves prevent blood from flowing backward in veins
a well-recognized sign of inflammation is adhesion of white
just as valves do in the heart, and represent another adaptation
blood cells to the postcapillary venule endothelium, followed
to compensate for low venous pressure. They are formed from
by their migration through the wall into the inflamed tissue.
folds of the tunica intima and resemble the semilunar valves of
Larger venules have a thin tunica media (one or two layers of
the heart ( p. 712) (see Figure 19.2). Venous valves are most
smooth muscle cells) and a thin tunica externa as well.
abundant in the veins of the limbs, where gravity opposes the
upward flow of blood. They are usually absent in veins of the
Veins thoracic and abdominal body cavities.
Venules join to form veins. Veins usually have three distinct tunics, The effectiveness of venous valves is demonstrated by this
but their walls are always thinner and their lumens larger than simple experiment: Hang one hand by your side until the blood
those of corresponding arteries (see Figure 19.2 and Table 19.1). vessels on its dorsal aspect distend with blood. Next place
Consequently, in histological preparations, veins are usually col- two fingertips against one of the distended veins, and pressing
lapsed and their lumens appear slitlike. firmly, move the superior finger proximally along the vein and
There is relatively little smooth muscle or elastin in the then release that finger. The vein will remain collapsed (flat)
tunica media, which is poorly developed and tends to be thin despite the pull of gravity. Finally, remove your distal fingertip
even in the largest veins. The tunica externa is the heaviest and watch the vein refill with blood.
wall layer. Consisting of thick longitudinal bundles of collagen
fibers and elastic networks, it is often several times thicker HOMEOSTATIC
than the tunica media. In the largest veins—the venae cavae, CLINICAL
IMBALANCE 19.1
which return blood directly to the heart—longitudinal bands of
Varicose veins are veins that are tortuous and dilated because
smooth muscle make the tunica externa even thicker.
of incompetent (leaky) valves. More than 15% of adults suffer
With their large lumens and thin walls, veins can accommo-
from varicose veins, usually in the lower limbs.
date a fairly large blood volume. Veins are called capacitance
Several factors contribute, including heredity and conditions
vessels and blood reservoirs because they can hold up to 65%
that hinder venous return, such as prolonged standing in one
of the body’s blood supply at any time (Figure 19.6). Even so,
position, obesity, or pregnancy. Both the “potbelly” of an over-
these distensible vessels are usually not filled to capacity.
weight person and the enlarged uterus of a pregnant woman
The walls of veins can be much thinner than arterial walls
exert downward pressure on vessels of the groin, restricting
without danger of bursting because the blood pressure in veins
return of blood to the heart. Consequently,
blood pools in the lower limbs, and with
time, the valves weaken and the venous
Pulmonary blood walls stretch. Superficial veins, which 19
17
vessels 12%
Systemic arteries Pulmonary blood vessels supply the lungs.
receive little support from surrounding
and arterioles 15% tissues, are especially susceptible.
Elevated venous pressure can also
cause varicose veins. For example, strain-
ing to deliver a baby or have a bowel
Heart 8%
movement raises intra-abdominal pres-
Capillaries 5% sure, preventing blood from draining
from anal veins. The resulting varicosities
in the anal veins are called hemorrhoids
Systemic veins
and venules 60% (hem9ŏ-roidz).
Systemic veins:
• Supply all of the body except the lungs
• Are distensible Venous Sinuses
• Contain a large proportion of the
blood volume and so are called Venous sinuses are highly specialized,
capacitance vessels or blood reservoirs flattened veins with extremely thin walls
composed only of endothelium. They are
Figure 19.6 Relative proportion of blood volume throughout the cardiovascular supported by the tissues that surround
system. them, rather than by any additional tunics.
(Do not confuse venous sinuses with the
M19_MARI0853_11_GE_C19.indd 745 03/07/18 2:05 PM

air-filled spaces in bones, the paranasal sinuses.) Examples of Like scaling a mountain, tackling blood pressure regulation
venous sinuses include the coronary sinus of the heart and the and other topics of cardiovascular physiology is challenging
dural venous sinuses of the brain. The dural venous sinuses, while you’re doing it, and exhilarating when you succeed. Let’s
which receive cerebrospinal fluid and blood draining from the begin the climb.
brain, are reinforced by the tough dura mater that covers the To sustain life, blood must be kept circulating. By now, you
brain surface ( p. 496). are aware that the heart is the pump, the arteries are pressure
reservoirs and conduits, the arterioles are resistance vessels that
Check Your Understanding control distribution, the capillaries are exchange sites, and the
5. What is the function of venous valves? What forms the valves? veins are conduits and blood reservoirs. Now for the dynamics
6. In the systemic circuit, which contains more blood—arteries or of this system.
veins—or is it the same?
19.6 Blood flows from high to low
19.5 Anastomoses are special pressure against resistance
Learning Outcome
interconnections between
NN Define blood flow, blood pressure, and resistance, and
blood vessels explain the relationships between these factors.
Learning Outcome First we need to define three physiologically important terms—
NN Explain the importance of vascular anastomoses. blood flow, blood pressure, and resistance—and examine how
these factors relate to the physiology of blood circulation.
Blood vessels form special interconnections called vascular
anastomoses (ah-nas0to-mo9sēz; “coming together”). Most
organs receive blood from more than one arterial branch, and Definition of Terms
arteries supplying the same territory often merge, forming arte- Blood Flow
rial anastomoses. These anastomoses provide alternate path- Blood flow is the volume of blood flowing through a vessel,
ways, called collateral channels, for blood to reach a given an organ, or the entire circulation in a given period (ml/min). If
body region. If one branch is cut or blocked by a clot, the col- we consider the entire vascular system, blood flow is equiva-
lateral channel can often provide sufficient blood to the area. lent to cardiac output (CO), and under resting conditions, it is
Arterial anastomoses occur around joints, where active relatively constant. At any given moment, however, blood flow
movement may hinder blood flow through one channel. They through individual body organs may vary widely according to
are also common in abdominal organs, the heart, and the brain their immediate needs.
(for example, the cerebral arterial circle in Figure 19.24c on
p. 773). Arteries that supply the retina, kidneys, and spleen Blood Pressure (BP)
either do not anastomose or have a poorly developed collateral
Blood pressure (BP), the force per unit area exerted on a ves-
circulation. If their blood flow is interrupted, cells supplied by
sel wall by the contained blood, is expressed in millimeters of
such vessels die.
mercury (mm Hg). For example, a blood pressure of 120 mm
19
17 The metarteriole–thoroughfare channel shunts of some cap-
Hg is equal to the pressure exerted by a column of mercury
illary beds that connect arterioles and venules are examples of
120 mm high.
arteriovenous anastomoses. Veins interconnect much more
Unless stated otherwise, the term blood pressure means
freely than arteries, and venous anastomoses are common.
systemic arterial blood pressure in the largest arteries near
(You may be able to see venous anastomoses through the skin
the heart. The hydrostatic pressure gradient—the differences
on the dorsum of your hand.) Because venous anastomoses are
in blood pressure within the vascular system—provides the
abundant, an occluded vein rarely blocks blood flow or leads
driving force that keeps blood moving, always from an area of
to tissue death.
higher pressure to an area of lower pressure, through the body.
Resistance
7. Which have more anastomoses, arteries or veins?
Resistance is opposition to f low and is a measure of the
amount of friction blood encounters as it passes through the
vessels. Because most friction is encountered in the peripheral
PART 2 (systemic) circulation, well away from the heart, we generally
use the term total peripheral resistance (TPR).
PHYSIOLOGY OF CIRCULATION There are three important sources of resistance: blood vis-
cosity, vessel length, and vessel diameter. You already know
Have you ever climbed a mountain? Well, get ready to climb a
more about these sources of resistance than you think you do. If
metaphorical mountain as you learn about circulatory dynamics.
M19_MARI0853_11_GE_C19.indd 746 03/07/18 2:05 PM

you wanted to drink this thick little to total peripheral resistance. Instead, the small-diameter
(viscous) milkshake, would arterioles, which can enlarge or constrict in response to neu-
you pick the short, wide straw ral and chemical controls, are the major determinants of total
or the long, narrow straw? We peripheral resistance.
bet you picked the short, wide When blood encounters either an abrupt change in vessel
straw. Let’s see why that makes diameter or rough or protruding areas of the tube wall (such
sense. as the fatty plaques of atherosclerosis), the smooth laminar
blood flow is replaced by turbulent flow, that is, irregular fluid
Blood Viscosity The inter- motion where blood from the different laminae (different lay-
nal resistance to f low that ers of the tube’s cross section) mixes. Turbulence dramatically
exists in all fluids is viscosity increases resistance.
(vis-kos9ĭ-te) and is related to
the thickness or “stickiness” of
a fluid. The greater the viscosity, the less easily molecules slide
Relationship between Flow, Pressure,
past one another and the more difficult it is to get and keep the and Resistance
fluid moving. Like our milkshake, blood is much more viscous Now that we have defined these terms, let’s summarize the rela-
than water. Because it contains formed elements and plasma pro- tionships between them.
teins, it flows more slowly under the same conditions. ●● Blood flow (F) is directly proportional to the difference in
Blood viscosity is fairly constant. However, conditions such blood pressure (ΔP) between two points in the circulation, that
as polycythemia (excessive numbers of red blood cells) can is, the blood pressure, or hydrostatic pressure, gradient. Con-
increase blood viscosity and so resistance increases. On the sequently, when ΔP increases, blood flow increases, and when
other hand, if the red blood cell count is low, as in some anemias, ΔP decreases, blood flow declines.
blood is less viscous and total peripheral resistance declines.
●● Blood flow is inversely proportional to the total periph-
Total Blood Vessel Length The relationship between total eral resistance (TPR) in the systemic circulation; if TPR
blood vessel length and resistance is straightforward: the longer increases, blood flow decreases.
the vessel, the greater the resistance. (That’s why a shorter We can express these relationships by the formula
straw is easier to drink from.) For example, an infant’s blood
vessels lengthen as he or she grows to adulthood, and so both ∆P
F=
total peripheral resistance and blood pressure increase. TPR
Blood Vessel Diameter The relationship between blood vessel Of these two factors influencing blood flow, TPR is far more
diameter and resistance is also straightforward: the smaller the important than ΔP in influencing local blood flow because TPR
diameter, the greater the resistance. (That’s why a wider straw can easily be changed by altering blood vessel diameter. For
is easier to drink from.) example, when the arterioles serving a particular tissue dilate
Blood viscosity and vessel length are normally unchanging (decreasing the resistance), blood flow to that tissue increases,
in the short term, and so the influence of these factors can be even though the systemic pressure is unchanged or may actually
considered constant. However, blood vessel diameter changes be falling.
frequently and significantly alters total peripheral resistance. 19
17
How so? The answer lies in principles of fluid flow. Fluid Check Your Understanding
close to the wall of a tube or channel is slowed by friction as 8. List three factors that determine resistance in a vessel. Which
it passes along the wall, whereas fluid in the center of the tube of these factors is physiologically most important?
flows more freely and faster. You can verify this by watching 9. APPLY Suppose vasoconstriction decreases the diameter of
the flow of water in a river. Water close to the bank hardly a vessel to one-third its size. What happens to the rate of
seems to move, while that in the middle of the river flows flow through that vessel? Calculate the expected size of the
quite rapidly. change.
In a tube of a given size, the relative speed and position of fluid 10. APPLY Here are three blood vessels. Assuming that the
in the different regions of the tube’s cross section remain constant, difference in pressure along the vessel length is the same for
a phenomenon called laminar flow or streamlining. The smaller each, which would have the greatest blood flow? The least
the tube, the greater the friction, because relatively more of the blood flow? Explain.
fluid contacts the tube wall, where its movement is impeded.
Resistance varies inversely with the fourth power of the ves- (a)
sel radius (one-half the diameter). This means, for example, (b)
that if the radius of a vessel doubles, the resistance drops to
one-sixteenth of its original value (r4 5 2 3 2 3 2 3 2 5 16
and 1/r4 5 1/16). For this reason, the large arteries close to the (c)
heart, which do not change dramatically in diameter, contribute
M19_MARI0853_11_GE_C19.indd 747 03/07/18 2:05 PM

aorta as aortic pressure reaches its peak. Indeed, if the aorta

Blood pressure decreases as
19.7 were opened during this period, blood would spurt upward 5 or
blood flows from arteries through 6 feet! This pressure peak generated by ventricular contraction
is called the systolic pressure (sis-tah9lik) and averages 120
capillaries and into veins mm Hg in healthy adults. Blood moves forward into the arterial
Learning Outcome bed because the pressure in the aorta is higher than the pressure
in the more distal vessels.
NN Describe how blood pressure differs in the arteries,
capillaries, and veins.
During diastole, the aortic valve closes, preventing blood
from flowing back into the heart. The walls of the aorta (and
Any fluid driven by a pump through a circuit of closed chan- other elastic arteries) recoil, maintaining sufficient pressure to
nels operates under pressure, and the nearer the fluid is to the keep the blood flowing forward into the smaller vessels. During
pump, the greater the pressure exerted on the fluid. Blood flow this time, aortic pressure drops to its lowest level (approxi-
in blood vessels is no exception, and blood flows through the mately 70 to 80 mm Hg in healthy adults). This is called the
blood vessels along a pressure gradient, always moving from diastolic pressure (di-as-tah9lik). You can picture the elastic
higher- to lower-pressure areas. Fundamentally, the pumping arteries as pressure reservoirs that operate as auxiliary pumps.
action of the heart generates blood flow. Pressure results when They keep blood circulating throughout the period of diastole,
flow is opposed by resistance. when the heart is relaxing. Essentially, the volume and energy of
As illustrated in Figure 19.7, systemic blood pressure is blood stored in the elastic arteries during systole are given back
highest in the aorta and declines throughout the pathway to during diastole.
finally reach 0 mm Hg in the right atrium. The steepest drop in The difference between the systolic and diastolic pressures is
blood pressure occurs in the arterioles, which offer the greatest called the pulse pressure. It is felt as a throbbing pulsation in an
resistance to blood flow. However, as long as a pressure gradi- artery (a pulse) during systole as ventricular contraction forces
ent exists, no matter how small, blood continues to flow until it blood into the elastic arteries and expands them. Increased
completes the circuit back to the heart. stroke volume and faster blood ejection from the heart (a result
of increased contractility) raise pulse pressure temporarily. Ath-
Arterial Blood Pressure erosclerosis chronically increases pulse pressure because the
Arterial blood pressure reflects two factors: (1) how much the elastic arteries become less stretchy.
elastic arteries close to the heart can stretch (their compliance Because aortic pressure fluctuates up and down with each
or distensibility) and (2) the volume of blood forced into them heartbeat, the important pressure to consider is the mean arte-
at any time. If the amounts of blood entering and leaving the rial pressure (MAP)—the pressure that propels the blood to
elastic arteries in a given period were equal, arterial pressure the tissues. Diastole usually lasts longer than systole, so MAP
would be constant. Instead, as Figure 19.7 reveals, blood pres- is not simply the value halfway between systolic and diastolic
sure is pulsatile—it rises and falls in a regular fashion—in the pressures. Instead, it is roughly equal to the diastolic pressure
elastic arteries near the heart. plus one-third of the pulse pressure.
As the left ventricle contracts and expels blood into the aorta, pulse pressure
it imparts kinetic energy to the blood, which stretches the elastic MAP = diastolic pressure +
3
19
17 For a person with a systolic blood pressure of 120 mm Hg
and a diastolic pressure of 80 mm Hg:
40 mm Hg
120 MAP = 80 mm Hg + = 93 mm Hg
3
Blood pressure (mm Hg)
Systolic pressure
100
Mean pressure MAP and pulse pressure both decline with increasing dis-
80 tance from the heart. The MAP loses ground to the never-
ending friction between the blood and the vessel walls, and the
60 pulse pressure is gradually phased out in the less elastic muscu-
Diastolic lar arteries, where elastic rebound of the vessels ceases to occur.
40 pressure At the end of the arterial tree, blood flow is steady and the pulse
20 pressure has disappeared.
0 Clinical Monitoring of Circulatory Efficiency

Clinicians can assess the efficiency of a person’s circulation by
r ta
ies
ies
ae
e
le
in
iol
av
Ao
nu
Ve
ter
lar
measuring pulse and blood pressure. These values, along with

ter
ec
Ve
pil
Ar
measurements of respiratory rate and body temperature, are

Ar
Ca
na
Ve
referred to collectively as the body’s vital signs. Let’s examine

Figure 19.7 Blood pressure in various blood vessels of the how vital signs are determined or measured.
systemic circulation.
M19_MARI0853_11_GE_C19.indd 748 03/07/18 2:05 PM

1. Wrap the blood pressure cuff, or sphygmomanometer
Superficial temporal artery (sfig0mo-mah-nom9ĕ-ter; sphygmo 5 pulse), snugly around
the person’s arm just superior to the elbow.
2. Inflate the cuff until the cuff pressure exceeds systolic pres-
Facial artery
sure. At this point, blood flow into the arm stops and a bra-
chial pulse cannot be felt or heard.
Common carotid artery 3. Reduce the cuff pressure gradually and listen (auscultate)
with a stethoscope for sounds in the brachial artery.
The pressure read when the first soft tapping sounds are
Brachial artery
heard (the first point at which a small amount of blood is spurt-
ing through the constricted artery) is systolic pressure. As the
cuff pressure is reduced further, these sounds, called the sounds
Radial artery
of Korotkoff, become louder and more distinct. However, when
the artery is no longer constricted and blood flows freely, the
Femoral artery sounds can no longer be heard. The pressure at which the
sounds disappear is the diastolic pressure.
Popliteal artery
Capillary Blood Pressure
As Figure 19.7 shows, by the time blood reaches the capillaries,
Posterior tibial blood pressure has dropped to approximately 35 mm Hg and by
artery
the end of the capillary beds is only around 17 mm Hg. Such
low capillary pressures are desirable because:
Dorsalis pedis
artery
●● Capillaries are fragile and high pressures would rupture them.
●● Most capillaries are extremely permeable and so even the low
Figure 19.8 Body sites where the pulse is most easily capillary pressure can force solute-containing fluids (filtrate)
palpated. (We discuss the specific arteries indicated on out of the bloodstream into the interstitial space.
pp. 770–781.)
As we describe later in this chapter, these fluid flows are
important for continuously refreshing the interstitial fluid.
Taking a Pulse You can feel a pulse in any artery that lies
close to the body surface by compressing the artery against Venous Blood Pressure
firm tissue, and this provides an easy way to count heart rate. Unlike arterial pressure, which pulsates with each contrac-
Because it is so accessible, the point where the radial artery tion of the left ventricle, venous blood pressure is steady and
surfaces at the wrist, the radial pulse, is routinely used to take a changes very little during the cardiac cycle. The pressure gradi-
pulse measurement, but there are several other clinically impor- ent in the veins, from venules to the termini of the venae cavae,
tant arterial pulse points (Figure 19.8). is only about 15 mm Hg (that from the aorta to the ends of the 19
17
These pulse points are also called pressure points because arterioles is about 60 mm Hg).
they are compressed to stop blood flow into distal tissues dur- The difference in pressure between an artery and a vein
ing hemorrhage. For example, if you seriously lacerate your becomes very clear when the vessels are cut. If a vein is cut,
hand, you can slow or stop the bleeding by compressing your the blood flows evenly from the wound, but a lacerated artery
radial or brachial artery. spurts blood. The very low pressure in the venous system
Monitoring pulse rate is an easy way to assess the effects reflects the cumulative effects of total peripheral resistance,
of activity, postural changes, and emotions on heart rate. For which dissipates most of the energy of blood pressure (as heat)
example, the pulse of a healthy man may be around 66 beats during each circuit.
per minute when he is lying down, 70 when he sits up, and 80 Despite the structural modifications of veins (large lumens
if he suddenly stands. During vigorous exercise or emotional and valves), venous pressure is normally too low to promote
upset, pulse rates between 140 and 180 are not unusual because adequate venous return. For this reason, three functional adap-
of sympathetic nervous system effects on the heart. tations are critically important to venous return:
Measuring Blood Pressure Most often, you measure systemic ●● The muscular pump. The muscular pump consists of skel-
arterial blood pressure indirectly in the brachial artery of the etal muscle activity. As the skeletal muscles surrounding the
arm by the auscultatory method (aw-skul9tah-to0re). The steps deep veins contract and relax, they squeeze or “milk” blood
of this procedure are:
M19_MARI0853_11_GE_C19.indd 749 05/07/18 5:20 PM


11. Use Figure 19.7 on p. 748 to answer the following questions:
(a) In which type of vessel is the largest pressure drop?
Venous valve (b) Which type of vessel is the first in which there is no difference
(open) in pressure between systole and diastole? Why is this a good
thing? (c) How much pressure remains at the start of the
venules to get blood back to the heart? What else helps blood
Contracted return to the heart?
skeletal
12. APPLY Cole has a systolic pressure of 140 and a diastolic
muscle
pressure of 80 mm Hg. What is his mean arterial pressure? His
pulse pressure?
Venous valve
(closed) Blood pressure is regulated by
19.8
short- and long-term controls
Learning Outcomes
Vein
NN List and explain the factors that influence blood pressure,
and describe how blood pressure is regulated.
NN Define hypertension. Describe its manifestations and
consequences.
NN Define circulatory shock. List several possible causes.
Direction of
blood flow Maintaining a steady flow of blood from the heart to the toes
is vital for organs to function properly. In fact, making sure a
person jumping out of bed in the morning does not keel over
from inadequate blood flow to the brain requires the finely
Figure 19.9 The muscular pump. When contracting skeletal
tuned cooperation of the heart, blood vessels, and kidneys—all
muscles press against a vein, they force open the valves proximal supervised by the brain.
to the area of contraction and blood is propelled toward the heart. Maintaining blood pressure is critical for cardiovascular sys-
Backflowing blood closes the valves distal to the area of contraction. tem homeostasis. Its regulation involves three key variables:
●● Cardiac output
●● Total peripheral resistance
toward the heart, and once blood passes each successive valve,
●● Blood volume
it cannot flow back (Figure 19.9). People who earn their liv-
ing in “standing professions,” such as hairdressers, often have To see why these are the central variables, we use the for-
swollen ankles because blood pools in their feet and legs. mula about blood flow presented on p. 747. In the cardiovas-
19
17
Indeed, standing for prolonged periods may cause fainting cular system, flow (F) is cardiac output (CO)—the blood flow
because skeletal muscle inactivity reduces venous return. of the entire circulation. P is blood pressure (MAP), and TPR
●● The respiratory pump. The respiratory pump moves is the total peripheral resistance (resistance of the blood vessels
blood up toward the heart as pressure changes in the ven- in the systemic circulation). If we rearrange the formula for blood
tral body cavity during breathing. As we inhale, abdominal flow, we can see how CO and TPR relate to blood pressure:
pressure increases, squeezing local veins and forcing blood F = ∆P>TPR or CO = ∆P>TPR or ∆P = CO × TPR
toward the heart. At the same time, the pressure in the chest
decreases, allowing thoracic veins to expand and speeding As you can see, blood pressure varies directly with CO and
blood entry into the right atrium. TPR. Anything that increases cardiac output or total peripheral
resistance increases blood pressure. Blood pressure also varies
●● Sympathetic venoconstriction. Sympathetic venoconstric-
directly with blood volume because CO depends on blood vol-
tion reduces the volume of blood in the veins—the capaci-
ume (the heart can’t pump out what doesn’t enter its chambers).
tance vessels. As the layer of smooth muscle around the
From Chapter 18 ( Figure 18.20, p. 729), you know that
veins constricts under sympathetic control, venous volume
CO is equal to stroke volume (ml/beat) times heart rate (beats/
is reduced and blood is pushed toward the heart.
min), so anything that increases these two variables will also
All three of these functional adaptations increase venous increase blood pressure. During stress, for example, the cardio-
return, which increases stroke volume (by the Frank-Starling acceleratory center activates the sympathetic nervous system,
mechanism, p. 729) and therefore increases cardiac output. which increases both heart rate (by acting on the SA node) and
M19_MARI0853_11_GE_C19.indd 750 03/07/18 2:05 PM

Stroke volume Heart rate Diameter of Blood viscosity Blood vessel

blood vessels length
Cardiac output Total peripheral resistance
Mean arterial pressure (MAP)
Figure 19.10 Major factors that increase MAP. In addition, cardiac output increases as
blood volume increases (not shown).
stroke volume (by enhancing cardiac muscle contractility). The changes in arterial pressure and stretch) and associated affer-
resulting increase in CO increases MAP. ent fibers. These reflexes are integrated in the cardiovascular
We also know that total peripheral resistance is determined center of the medulla, and their output travels via autonomic
by three variables, the most important of which is blood ves- fibers to the heart and vascular smooth muscle. Occasionally,
sel diameter ( pp. 746–747). Figure 19.10 summarizes the inputs from chemoreceptors (receptors that respond to changes
relationships between the factors controlling CO and resistance. in blood levels of carbon dioxide, H+, and oxygen) and higher
Keep these relationships in mind as you read through the sec- brain centers also influence the neural control mechanism.
tions that follow, because each blood pressure regulation mech-
anism acts on one or more of these variables. Role of the Cardiovascular Center
Also be aware that things aren’t quite that simple in real life. Several clusters of neurons in the medulla oblongata act
A change in any variable that threatens blood pressure homeo- together to integrate blood pressure control by altering cardiac
stasis is usually compensated for by changes in the other vari- output and blood vessel diameter. This cardiovascular center
ables so that a constant blood pressure is maintained. consists of the cardiac centers (the cardioacceleratory and car-
We will now explore two classes of mechanisms that regulate dioinhibitory centers discussed in Chapter 18, p. 721) and the
blood pressure. Short-term regulation by the nervous system vasomotor center that controls the diameter of blood vessels.
and bloodborne hormones alters blood pressure by changing The vasomotor center transmits impulses at a fairly steady
total peripheral resistance and CO. Long-term regulation alters rate along sympathetic efferents called vasomotor fibers.
blood volume via the kidneys. Figure 19.13 (p. 755) summarizes These fibers exit from the T1 through L2 levels of the spinal
the influence of nearly all of the important factors. cord and innervate the smooth muscle of blood vessels, mainly
arterioles. As a result, the arterioles are almost always in a state
Short-Term Regulation: Neural Controls of moderate constriction, called vasomotor tone ( p. 577).
Neural controls alter both cardiac output and total peripheral The degree of vasomotor tone varies from organ to organ.
19
17
resistance. We discussed neural control of cardiac output in Generally, arterioles of the skin and digestive viscera receive
Chapter 18, so we will focus on total peripheral resistance here. vasomotor impulses more frequently and tend to be more
Neural controls of total peripheral resistance are directed at two strongly constricted than those of skeletal muscles. Any
main goals: increase in sympathetic activity produces generalized vaso-
●● Maintaining adequate MAP by altering blood vessel diam- constriction and raises blood pressure. Decreased sympathetic
eter on a moment-to-moment basis. (Remember, very small activity allows the vascular smooth muscle to relax somewhat
changes in blood vessel diameter cause substantial changes and lowers blood pressure to basal levels.
in total peripheral resistance, and so in systemic blood pres- Cardiovascular center activity is modified by inputs from
sure.) Under conditions of low blood volume, all vessels baroreceptors, chemoreceptors, and higher brain centers. Let’s
except those supplying the heart and brain are constricted to take a look.
allow as much blood as possible to flow to those two vital
Baroreceptor Reflexes
organs.
●● Altering blood distribution to respond to specific demands When arterial blood pressure rises, it activates baroreceptors.
of various organs. For example, during exercise blood is These stretch receptors are located in the carotid sinuses (dila-
shunted temporarily from the digestive organs to the skeletal tions in the internal carotid arteries, which provide the major
muscles. blood supply to the brain), in the aortic arch, and in the walls
of nearly every large artery of the neck and thorax. When
Most neural controls operate via reflex arcs involving baro- stretched, baroreceptors send a rapid stream of impulses to the
receptors (pressure-sensitive mechanoreceptors that respond to
M19_MARI0853_11_GE_C19.indd 751 03/07/18 2:05 PM

3 Impulses from baroreceptors activity, reducing heart rate

stimulate cardioinhibitory center and contractile force. As
(and inhibit cardioacceleratory CO falls, so does MAP.
center) and inhibit vasomotor
center. In the opposite situation, a
4a Sympathetic decline in MAP initiates reflex
impulses to heart vasoconstriction and increases
cause HR,
contractility, and
cardiac output, bringing blood
CO. pressure back up. In this way,
total peripheral resistance and
2 Baroreceptors
in carotid sinuses
cardiac output are regulated in
and aortic arch tandem to minimize changes in
are stimulated. blood pressure.
Rapidly responding baro-
4b Rate of
vasomotor impulses
receptors protect the circula-
allows vasodilation, tion against short-term (acute)
IMB
AL
causing TPR. 5 CO and TPR changes in blood pressure. For
1 Stimulus: AN return blood example, blood pressure falls
CE
Blood pressure pressure to
(arterial blood
(particularly in the head) when
homeostatic range.
pressure rises above Homeostasis: Blood pressure in normal range you stand up after reclining.
normal range). Baroreceptors taking part in
1 Stimulus: the carotid sinus reflex pro-
Blood pressure tect the blood supply to your
IMB (arterial blood brain, whereas those activated
AL
AN pressure falls below
CE
normal range).
in the aortic reflex help main-
5 CO and TPR
return blood pressure
tain adequate blood pressure
to homeostatic range.
4b Vasomotor in your systemic circuit as a
fibers stimulate
vasoconstriction, whole. Failure of the barore-
causing TPR. ceptor reflex results in ortho-
static hypotension ( p. 581).
Baroreceptors are rela-
2 Baroreceptors tively ineffective in protecting
in carotid sinuses us against sustained pressure
and aortic arch changes, as evidenced by the
are inhibited.
fact that many people develop
chronic hypertension. In such
4a Sympathetic
impulses to heart cases, the baroreceptors are
cause HR, “reprogrammed” (adapt) to
19
17 contractility, and 3 Impulses from baroreceptors
activate cardioacceleratory center monitor pressure changes at a
CO.
(and inhibit cardioinhibitory center) higher set point.
and stimulate vasomotor center.
Chemoreceptor Reflexes
Figure 19.11 Baroreceptor reflexes that help maintain blood pressure homeostasis.
(CO 5 cardiac output; TPR 5 total peripheral resistance; HR 5 heart rate; BP 5 blood pressure) When the carbon dioxide lev-
els rise, or the pH falls, or oxy-
gen content of the blood drops
sharply, chemoreceptors in
cardiovascular center, inhibiting the vasomotor and cardio-
the aortic arch and large arteries of the neck transmit impulses
acceleratory centers and stimulating the cardioinhibitory center.
to the cardioacceleratory center, which then increases cardiac
The result is a decrease in blood pressure (Figure 19.11).
output. Chemoreceptors also activate the vasomotor center,
Two mechanisms bring this about:
which causes reflex vasoconstriction. The rise in blood pressure
●● Vasodilation. Decreased output from the vasomotor center that follows speeds the return of blood to the heart and lungs.
allows arterioles and veins to dilate. Arteriolar vasodilation The most prominent chemoreceptors are the carotid and
reduces total peripheral resistance, so MAP falls. Venodila- aortic bodies located close by the baroreceptors in the carotid
tion shifts blood to the venous reservoirs, which decreases sinuses and aortic arch. Chemoreceptors play a larger role in
venous return and cardiac output. regulating respiratory rate than blood pressure, so we consider
●● Decreased cardiac output. Impulses to the cardiac centers their function in Chapter 22.
inhibit sympathetic activity and stimulate parasympathetic
M19_MARI0853_11_GE_C19.indd 752 03/07/18 2:05 PM

Influence of Higher Brain Centers ●● Antidiuretic hormone (ADH). Produced by the hypothala-
Reflexes that regulate blood pressure are integrated in the mus, antidiuretic hormone (ADH, also called vasopressin)
medulla oblongata of the brain stem. Although the cerebral cor- stimulates the kidneys to conserve water ( p. 645). It is
tex and hypothalamus are not involved in routine controls of not usually important in short-term blood pressure regula-
blood pressure, these higher brain centers can modify arterial tion. However, when blood pressure falls to dangerously
pressure via relays to the medullary centers. low levels (as during severe hemorrhage), much more ADH
For example, the fight-or-flight response mediated by the is released and helps restore arterial pressure by causing
hypothalamus has profound effects on blood pressure. (Even intense vasoconstriction.
the simple act of speaking can make your blood pressure jump
if the person you are talking to makes you anxious.) The hypo- Long-Term Regulation: Renal Mechanisms
thalamus also mediates the redistribution of blood flow and Unlike short-term controls of blood pressure that alter total
other cardiovascular responses that occur during exercise and peripheral resistance and cardiac output, long-term controls
changes in body temperature. alter blood volume. Renal mechanisms mediate long-term regu-
lation by the kidneys.
Short-Term Regulation: Hormonal Controls Although baroreceptors respond to short-term changes in
Hormones also help regulate blood pressure, both in the short blood pressure, they quickly adapt to prolonged or chronic epi-
term via changes in total peripheral resistance and in the long sodes of high or low pressure. This is where the kidneys step in
term via changes in blood volume (Table 19.2). Paracrines to restore and maintain blood pressure homeostasis by regulat-
(local chemicals), on the other hand, primarily serve to match ing blood volume. Although blood volume varies with age, body
blood flow to the metabolic need of a particular tissue. In rare size, and sex, renal mechanisms usually keep it close to 5 L.
instances, massive release of paracrines can affect blood pres- As we noted earlier, blood volume is a major determinant
sure. We will discuss these paracrines later—here we will of cardiac output [via its influence on venous return, end dias-
examine the short-term effects of hormones. tolic volume (EDV), and stroke volume]. An increase in blood
●● Adrenal medulla hormones. During periods of stress, the volume is followed by a rise in blood pressure. Anything that
adrenal gland releases epinephrine and norepinephrine increases blood volume—such as excessive salt intake, which
(NE) to the blood. Both hormones enhance the sympathetic promotes water retention—raises MAP because of the greater
response by increasing cardiac output and promoting general- fluid load in the vascular tree.
ized vasoconstriction. By the same token, decreased blood volume translates to a fall
in blood pressure. Dehydration that occurs during vigorous exer-
●● Angiotensin II. When blood pressure or blood volume are
cise and blood loss are common causes of reduced blood volume.
low, the kidneys release renin. Renin acts as an enzyme, ulti-
A sudden drop in blood pressure often signals internal bleeding
mately generating angiotensin II (an0je-o-ten9sin), which
and blood volume too low to support normal circulation.
stimulates intense vasoconstriction, promoting a rapid rise in
However, these assertions—increased blood volume
systemic blood pressure. It also stimulates release of aldos-
increases BP and decreased blood volume decreases BP—do
terone and ADH, which act in long-term regulation of blood
not tell the whole story because we are dealing with a dynamic
pressure as described on pp. 754–755.
system. Increases in blood volume that raise blood pressure
●● Atrial natriuretic peptide (ANP). The atria of the heart produce also stimulate the kidneys to eliminate water, which reduces
the hormone atrial natriuretic peptide (ANP), which leads to 19
17
blood volume and consequently blood pressure. Likewise, fall-
a reduction in blood volume and blood pressure. As noted in ing blood volume triggers renal mechanisms that increase blood
Chapter 16 ( p. 656), ANP antagonizes aldosterone and prods volume and blood pressure. As you can see, blood pressure can
the kidneys to excrete more sodium and water from the body, be stabilized or maintained within normal limits only when
reducing blood volume. It also causes generalized vasodilation. blood volume is stable.
Table 19.2 Effects of Selected Hormones on Blood Pressure

HORMONE EFFECT ON BP VARIABLE AFFECTED SITE OF ACTION
Epinephrine and norepinephrine (NE) c c CO (HR and contractility) Heart (b1 receptors)
c Total peripheral resistance (vasoconstriction) Arterioles (a receptors)
Angiotensin II c c Total peripheral resistance (vasoconstriction) Arterioles
Antidiuretic hormone (ADH) c c Total peripheral resistance (vasoconstriction) Arterioles
c Blood volume (T water loss) Kidney tubule cells
Aldosterone c c Blood volume (T salt and water loss) Kidney tubule cells
Atrial natriuretic peptide (ANP) T T Blood volume (c salt and water loss) Kidney tubule cells
T Total peripheral resistance (vasodilation) Arterioles
M19_MARI0853_11_GE_C19.indd 753 03/07/18 2:06 PM

Direct renal mechanism Indirect renal mechanism (renin-angiotensin-aldosterone)
Initial stimulus
Arterial pressure Arterial pressure
Physiological response
Result
Inhibits baroreceptors
Sympathetic nervous
system activity
Filtration by kidneys Angiotensinogen
Renin release
from kidneys
Angiotensin I
Angiotensin converting
enzyme (ACE)
Angiotensin II
Urine formation
ADH release by Thirst via Vasoconstriction; total

Adrenal cortex
posterior pituitary hypothalamus peripheral resistance
Secretes
Aldosterone
Blood volume
Sodium reabsorption Water reabsorption Water intake
by kidneys by kidneys
Blood volume
19
17
Mean arterial pressure Mean arterial pressure
Figure 19.12 Direct and indirect (hormonal) mechanisms for renal control of blood
pressure. Low blood pressure also triggers other actions not shown here that increase BP:
additional mechanisms of renin release (described in Chapter 25) and short-term actions of the
sympathetic nervous system.
The kidneys act both directly and indirectly to regulate arte- reabsorb the filtrate rapidly enough, and more of it leaves the
rial pressure and provide the major long-term mechanisms of body in urine. As a result, blood volume and blood pressure fall.
blood pressure control. When blood pressure or blood volume is low, water is con-
served and returned to the bloodstream, and blood pressure
Direct Renal Mechanism rises (Figure 19.12). As blood volume goes, so goes the arte-
The direct renal mechanism alters blood volume independently rial blood pressure.
of hormones. When either blood volume or blood pressure rises,
the rate at which fluid filters from the bloodstream into the kid- Indirect Renal Mechanism
ney tubules speeds up. In such situations, the kidneys cannot The kidneys can also regulate blood pressure indirectly via the
renin-angiotensin-aldosterone mechanism. When arterial
M19_MARI0853_11_GE_C19.indd 754 03/07/18 2:06 PM

Activity of Release Fluid loss from Crisis stressors: Vasomotor tone; Dehydration, Body size
muscular of ANP hemorrhage, exercise, trauma, bloodborne high hematocrit
pump and excessive body chemicals
respiratory sweating temperature (epinephrine,
pump NE, ADH,
angiotensin II)
Conservation Blood volume Blood pH

of Na1 and Blood pressure O2
water by kidneys CO2
Blood Baroreceptors Chemoreceptors

volume
Venous Activation of vasomotor and cardio-

return acceleratory centers in brain stem
Diameter of Blood Blood vessel

Stroke Heart
blood vessels viscosity length
volume rate
Cardiac output Total peripheral resistance
Initial stimulus
Physiological response
Mean arterial pressure (MAP)
Result
Figure 19.13 Factors that increase MAP.
blood pressure declines, certain cells in the kidneys release the ●● It is a potent vasoconstrictor, increasing blood pressure by
enzyme renin into the blood. Renin enzymatically splits angio- increasing total peripheral resistance. 19
17
tensinogen, a plasma protein made by the liver, converting it to
angiotensin I. In turn, angiotensin converting enzyme (ACE) Summary of Blood Pressure Regulation
converts angiotensin I to angiotensin II. ACE is found in the
capillary endothelium in various body tissues, particularly the How do each of the different
lungs. mechanisms that we have just Complete an interactive
Angiotensin II acts in four ways to stabilize arterial blood explored act together to control tutorial: >
Study Area > Interactive
pressure and extracellular fluid volume (Figure 19.12). blood pressure? Figure 19.13 pro- Physiology
vides a summary of how mean arte-
●● It stimulates the adrenal cortex to secrete aldosterone, a hor- rial pressure is controlled in concert
mone that enhances renal reabsorption of sodium from the by short- and long-term mechanisms. Notice that the left part of
forming urine. As sodium moves back into the bloodstream, the figure (the factors that control cardiac output) builds upon
water follows, which conserves blood volume. In addition, what you learned in Chapter 18 ( Figure 18.20, p. 729).
angiotensin II directly stimulates sodium reabsorption by the The goal of blood pressure regulation is to keep blood pres-
kidneys. sure high enough to provide adequate tissue perfusion (blood
●● It prods the posterior pituitary to release ADH, which pro- flow), but not so high that blood vessels are damaged. Consider
motes more water reabsorption by the kidneys. the brain. If pressure is too low, then perfusion is inadequate
●● It triggers the sensation of thirst by activating the hypotha- and you lose consciousness. If pressure is too high, your fragile
lamic thirst center (see Chapter 26). This encourages water brain capillaries might rupture and you would have a stroke.
consumption, ultimately restoring blood volume and so Malfunction of blood pressure control is our next topic.
blood pressure.
M19_MARI0853_11_GE_C19.indd 755 03/07/18 2:06 PM

and interfere with the ability of endothelial cells to induce

Homeostatic Imbalances CLINICAL
vasodilation.
in Blood Pressure ●● Age. Hypertension usually appears after age 40.
Normal blood pressure for resting adults is a systolic pressure
●● Diabetes mellitus.
of less than 120 mm Hg and a diastolic pressure of less than
80 mm Hg. Transient elevations in blood pressure occur as normal ●● Stress. Particularly at risk are “hot reactors,” people whose
adaptations during changes in posture, physical exertion, emo- blood pressure zooms upward during every stressful event.
tional upset, and fever. Age, sex, weight, and race also affect ●● Smoking. Nicotine causes intense vasoconstriction not only
blood pressure. by directly stimulating postganglionic sympathetic neurons
but also by prompting release of large amounts of epineph-
Hypertension rine and NE. Chemicals in cigarette smoke also damage the
Chronically elevated blood pressure is called hypertension and tunica intima, interfering with its ability to chemically regu-
is characterized by a sustained increase in either systolic pres- late arteriolar diameter.
sure (above 140 mm Hg) or diastolic pressure (above 90 mm
Primary hypertension cannot be cured, but most cases can
Hg). The American Heart Association considers individuals to
be controlled. Improving diet, increasing exercise and losing
have prehypertension if their blood pressure values are elevated
weight, stopping smoking, managing stress, and taking anti
but not yet in the hypertension range. These individuals are
hypertensive drugs can all help. Drugs commonly used are
at higher risk for developing full-blown hypertension and are
diuretics, beta-blockers, calcium channel blockers, angiotensin
often advised to change their lifestyles to reduce their risk of
converting enzyme (ACE) inhibitors, and angiotensin II recep-
developing full-blown hypertension.
tor blockers. Inhibiting ACE or blocking receptors for angioten-
Chronic hypertension is a common and dangerous disease.
sin II suppresses the renin-angiotensin-aldosterone mechanism.
An estimated 30% of people over age 50 are hypertensive.
Although this “silent killer” is usually asymptomatic for the first Secondary Hypertension Secondary hypertension accounts
10 to 20 years, it slowly but surely strains the heart and damages for 10% of cases. It is due to identifiable conditions, for exam-
the arteries. Prolonged hypertension is the major cause of heart ple obstructed renal arteries, kidney disease, and endocrine
failure, vascular disease, renal failure, and stroke. The higher the disorders such as hyperthyroidism and Cushing’s syndrome.
pressure, the greater the risk for these serious problems. Treatment for secondary hypertension focuses on correcting the
Because the heart is forced to pump against greater resistance, problem that caused it.
it must work harder, and over time the myocardium enlarges.
When finally strained beyond its capacity, the heart weakens Hypotension
and its walls become flabby. Hypertension also ravages the In many cases, hypotension, or low blood pressure (below
blood vessels, accelerating the progress of atherosclerosis (see 90/60 mm Hg), simply reflects individual variations and is no
A Closer Look on p. 792). As the vessels become increasingly cause for concern. In fact, low blood pressure is often associ-
blocked, blood flow to the tissues becomes inadequate and vascu- ated with long life and an old age free of cardiovascular disease.
lar complications appear in the brain, heart, kidneys, and retinas Orthostatic hypotension is a temporary drop in blood pres-
of the eyes. sure resulting in dizziness (due to inadequate oxygen delivery
to the brain) when a person rises suddenly from a reclining or
Primary Hypertension Although hypertension and atheroscle-
19
17 sitting position. Elderly people are prone to orthostatic hypoten-
rosis are often linked, it is often difficult to blame hyperten-
sion because the aging sympathetic nervous system does not
sion on any distinct anatomical pathology. Indeed, about 90%
respond as quickly as it once did to postural changes. Blood
of hypertensive people have primary, or essential, hyperten-
pools briefly in the lower limbs, reducing blood pressure and
sion, for which no underlying cause has been identified. This is
consequently blood delivery to the brain. Changing position
because primary hypertension is due to a rich interplay between
slowly gives the nervous system time to adjust and usually pre-
your genes and a variety of environmental factors:
vents this problem.
●● Heredity. Children of hypertensive parents are twice as likely Occasionally, chronic hypotension is a sign of a serious
to develop hypertension as are children of normotensive par- underlying condition. Addison’s disease (inadequate adrenal
ents, and more blacks than whites are hypertensive. Many of cortex function), hypothyroidism, or severe malnutrition can
the factors listed here require a genetic predisposition, and the cause chronic hypotension.
course of the disease varies in different population groups. Hypotension is usually a concern only if it leads to inad-
●● Diet. Dietary factors that contribute to hypertension include equate blood flow to tissues. Acute hypotension is one of the
high intakes of salt (NaCl), saturated fat, and cholesterol, most important signs of circulatory shock.
and deficiencies in certain metal ions (K+, Ca2+, and Mg2+).
Circulatory Shock
●● Obesity. Obesity causes hypertension in a number of ways
Circulatory shock is any condition in which blood vessels are
that are not yet well understood. For example, adipocytes
inadequately filled and blood cannot circulate normally. Blood
release hormones that appear to increase sympathetic tone
M19_MARI0853_11_GE_C19.indd 756 03/07/18 2:06 PM

flow is inadequate to meet tissue needs. If circulatory shock
persists, cells die and organ damage follows. Intrinsic and extrinsic controls
19.9
Hypovolemic Shock The most common form of circulatory determine blood flow through tissues
shock is hypovolemic shock (hi0po-vo-le9mik; hypo 5 low, Learning Outcome
deficient; volemia 5 blood volume), which results from large-
NN Explain how blood flow through tissues is regulated in
scale blood or fluid loss, as might follow acute hemorrhage, general and in specific organs.
severe vomiting or diarrhea, or extensive burns. If blood volume
drops rapidly, heart rate increases in an attempt to correct the Blood f low through body tissues, or tissue perfusion, is
problem. A weak, “thready” (barely detectable) pulse is often involved in:
the first sign of hypovolemic shock. Intense vasoconstriction ●● Delivering oxygen and nutrients to tissue cells, and removing
also occurs, which shifts blood from the various blood reser- carbon dioxide and wastes
voirs into the major circulatory channels and enhances venous ●● Exchanging gases in the lungs
return.
Blood pressure is stable at first, but eventually drops if blood ●● Absorbing nutrients from the digestive tract
loss continues. A sharp drop in blood pressure is a serious, and ●● Forming urine in the kidneys
late, sign of hypovolemic shock. The key to managing hypo- The rate of blood f low to each tissue and organ is almost
volemic shock is to replace fluid volume as quickly as possible. exactly the right amount to provide for proper function—no
Vascular Shock In vascular shock, blood volume is normal, more, no less. This is achieved by intrinsic controls (autoreg-
but circulation is poor as a result of extreme vasodilation. A ulation) acting automatically on the smooth muscle of arteri-
huge drop in total peripheral resistance follows, as revealed by oles that feed any given tissue. We will examine these intrinsic
rapidly falling blood pressure. mechanisms in the next section.
There are three common types of vascular shock: First, let’s step back and look at the big picture. What do you
think would happen if all of the arterioles in your body dilated
●● Anaphylactic shock is a loss of vasomotor tone due to ana-
at once? Because there is only a finite amount of blood, blood
phylaxis, a systemic allergic reaction in which the massive
pressure would fall. Critical tissues, such as the brain, would be
release of the chemical messenger histamine triggers body-
deprived of the oxygen and nutrients they need and would stop
wide vasodilation.
functioning. Extrinsic controls keep this from happening by
●● Neurogenic shock results from failure of autonomic nervous acting on arteriolar smooth muscle to maintain blood pressure.
system regulation. The extrinsic controls act via the nerves (sympathetic nervous
●● Septic shock (septicemia) is due to severe systemic bacterial system) and hormones of the nervous and endocrine systems,
infection (bacterial toxins are notorious vasodilators). the two major control systems of the body. They reduce blood
flow to regions that need it the least, maintaining a constant
Cardiogenic Shock Cardiogenic shock, or pump failure, MAP and allowing intrinsic mechanisms to direct blood flow
occurs when the heart is so inefficient that it cannot sustain to where it is most needed.
adequate circulation. Its usual cause is myocardial damage, as A number of physiological processes are under both intrinsic
might follow numerous myocardial infarctions (heart attacks). and extrinsic control (Figure 19.14). (Note that these are con-
trol mechanisms, and not related in any way to the intrinsic and 19
17
Check Your Understanding extrinsic pathways of blood clotting.)
13. Describe the baroreceptor reflex changes that occur to maintain
blood pressure when you rise from a lying-down to a standing
position.
14. The kidneys play an important role in maintaining MAP by Intrinsic controls Extrinsic controls
influencing which variable? Explain how renal artery obstruction • Control is entirely from within • Control is from outside of
could cause secondary hypertension. the tissue or organ the tissue or organ
• Uses paracrines or properties • Uses nerves or hormones
15. APPLY Your neighbor, Bob, calls you because he thinks he of muscle tissue
is having an allergic reaction to a medication. You find Bob • Also known as autoregulation
on the verge of losing consciousness and having trouble or local control
breathing. When paramedics arrive, they note his blood
pressure is 63/38 and he has a rapid, thready pulse. Explain
Bob’s low blood pressure and rapid heart rate. Examples where both control mechanisms occur:
16. MAKE CONNECTIONS You have just learned that hypertension • Stroke volume in heart ( pp. 729–730)
can be treated with a variety of different drugs including • Arteriolar diameter (see Figure 19.16)
• Glomerular filtration in kidneys (see Chapter 25)
diuretics, beta-blockers, and calcium channel blockers. Using
your knowledge of the autonomic nervous system (Chapter 14), Figure 19.14 A quick summary of intrinsic versus extrinsic
smooth muscle (Chapter 9), and cardiac muscle (Chapter 18), control mechanisms.
explain how these drugs work to decrease blood pressure.
M19_MARI0853_11_GE_C19.indd 757 05/07/18 5:21 PM

You can compare blood flow autoregulation to water use in

750
your home. Whether you have several taps open or none, the
pressure in the main water pipe in the street remains relatively
constant, as it does in the even larger water lines closer to the
750 pumping station. Similarly, local conditions in the arterioles
feeding the capillary beds of an organ have little effect on pres-
Brain 750
12,500
sure in the muscular artery feeding that organ, or in the large
Heart 250 elastic arteries. The pumping station is, of course, the heart. As
long as the water company (circulatory feedback mechanisms)
Skeletal 1200 maintains a relatively constant water pressure (MAP), local
muscles demand regulates the amount of fluid (blood) delivered to vari-
ous areas.
Skin 500
Organs regulate their own blood flows by varying the resis-
tance of their arterioles. These intrinsic control mechanisms
Kidneys 1100 may be classed as metabolic (chemical) or myogenic (physical).
Generally, both metabolic and myogenic factors determine the
final autoregulatory response of a tissue. For example, reactive
Abdomen 1400 hyperemia (hi0per-e9me-ah) refers to the dramatically increased
1900
blood flow into a tissue that occurs after the blood supply to the
area has been temporarily blocked. It results both from the myo-
Other 600 genic response and from the metabolic wastes that accumulated
600 during occlusion. Figure 19.16 summarizes the various intrin-
Total blood
flow at rest sic (local) and extrinsic controls of arteriolar diameter.
5800 ml/min 600
400
Metabolic Controls
When blood flow is too low to meet a tissue’s metabolic needs,
Total blood flow during
strenuous exercise oxygen levels decline and metabolic products (which act as par-
17,500 ml/min acrines) accumulate. These changes serve as stimuli that lead to
automatic increases in tissue blood flow.
Figure 19.15 Distribution of blood flow at rest and during The metabolic factors that regulate blood flow are low oxy-
strenuous exercise.
gen levels, and increases in H+ (from CO2 and lactic acid), K+,
adenosine, and prostaglandins. The relative importance of these
factors is not clear. Many of them act directly to relax vascular
The redistribution of blood during exercise provides an smooth muscle, but some may act by causing vascular endothe-
example of how intrinsic and extrinsic controls work together lial cells to release nitric oxide.
(Figure 19.15). When the body is at rest, the brain receives Nitric oxide (NO) is a powerful vasodilator, but its effects
about 13% of total blood flow, the heart 4%, kidneys 20%, are very brief because it is quickly destroyed. Even so, NO is
19
17 and abdominal organs 24%. Skeletal muscles, which make the major player in controlling local vasodilation, often over-
up almost half of body mass, normally receive about 20% of riding sympathetic vasoconstriction when tissues need more
total blood flow. During exercise, however, nearly all of the blood flow.
increased cardiac output flushes into the skeletal muscles as The endothelium also releases potent vasoconstrictors,
intrinsic autoregulatory controls dilate skeletal muscle arteri- including the family of peptides called endothelins, which are
oles. To maintain blood pressure in spite of the widespread dila- among the most potent vasoconstrictors known. Normally, NO
tion of arterioles in skeletal muscle, the extrinsic controls act to and endothelin released from endothelial cells are in a dynamic
decrease blood flow to the kidneys and digestive organs. balance, but this balance tips in favor of NO when blood flow
is too low for metabolic needs.
Autoregulation: Intrinsic (Local) The net result of metabolically controlled autoregulation is
Regulation of Blood Flow immediate vasodilation of the arterioles serving the capillary
As our activities change throughout the day, how does each beds of the “needy” tissues. Blood flow to the area rises tem-
organ or tissue manage to get the blood flow it needs? The porarily, allowing blood to surge through the capillaries and
answer is autoregulation. Local conditions regulate blood become available to the tissue cells.
flow independent of control by nerves or hormones. Changes Inflammatory chemicals released in injury, infection, or
in blood flow through individual organs are controlled intrinsi- allergic reactions also cause local vasodilation. Inflammatory
cally by modifying the diameter of local arterioles feeding the vasodilation helps the defense mechanisms clear microorgan-
capillaries. isms and toxins from the area, and promotes healing.
M19_MARI0853_11_GE_C19.indd 758 03/07/18 2:06 PM

Vasodilators
Metabolic Neural
O2 Sympathetic tone
CO2
H1 Hormonal
K1
• Atrial natriuretic peptide
• Prostaglandins
• Adenosine
• Nitric oxide
Intrinsic controls Extrinsic controls

(autoregulation) • Neural or hormonal controls
Vasoconstrictors • Maintain mean arterial pressure
• Metabolic or myogenic controls
(MAP)
• Distribute blood flow to individual
• Redistribute blood during exercise
organs and tissues as needed
and thermoregulation
Myogenic Neural
• Stretch Sympathetic tone
Metabolic Hormonal
• Endothelins • Angiotensin II
• Antidiuretic hormone
• Epinephrine
• Norepinephrine
Figure 19.16 Intrinsic and extrinsic control of arteriolar smooth muscle in the
systemic circulation. Controls are listed in the boxes below the arterioles. Epinephrine
and norepinephrine constrict arteriolar smooth muscle by acting at a-adrenergic receptors.
b-adrenergic receptors (causing vasodilation) are present in arterioles supplying skeletal and
heart muscle, but their physiological relevance is minimal. 19
17
Myogenic Controls Long-Term Autoregulation

Fluctuations in systemic blood pressure would cause problems If a tissue needs more nutrients than short-term autoregulatory
for individual organs were it not for the myogenic responses mechanisms can supply, a long-term autoregulatory mechanism
(myo 5 muscle, gen 5 origin) of vascular smooth muscle. Inad- may develop over weeks or months to enrich local blood flow still
equate blood perfusion through an organ is quickly followed more. The number of blood vessels in the region increases, and
by a decline in the organ’s metabolic rate and, if prolonged, existing vessels enlarge. This phenomenon, called angiogenesis,
organ death. Likewise, excessively high arterial pressure can be is particularly common in the heart when a coronary vessel is
dangerous because it may rupture more fragile blood vessels. partially occluded. It occurs throughout the body in people who
Fortunately, vascular smooth muscle prevents these problems live in high-altitude areas, where the air contains less oxygen.
by responding directly to passive stretch (caused by increased
intravascular pressure) with increased tone, which resists the Blood Flow in Special Areas
stretch and causes vasoconstriction. In contrast, reduced stretch
Each organ has special requirements and functions that are
(due to decreased intravascular pressure) results in vasodila-
revealed in its pattern of autoregulation. Autoregulation in the
tion and increases blood flow into the tissue. These myogenic
brain, heart, and kidneys is extraordinarily efficient, maintain-
responses keep tissue perfusion fairly constant despite most
ing adequate perfusion even when MAP fluctuates.
variations in systemic pressure.
M19_MARI0853_11_GE_C19.indd 759 03/07/18 2:06 PM

Skeletal Muscles can increase tenfold or more during physical activity, as you
Blood flow in skeletal muscle varies with fiber type and muscle saw in Figure 19.15, and virtually all capillaries in the active
activity. Generally speaking, capillary density and blood flow are muscles open to accommodate the increased flow.*
greater in red (slow oxidative) fibers than in white (fast glycolytic)
The Brain
fibers. Resting skeletal muscles receive about 1 L of blood per
minute, and only about 25% of their capillaries are open. During Blood flow to the brain averages 750 ml/min and is maintained
rest, myogenic and general neural mechanisms predominate. at a relatively constant level. Constant cerebral blood flow is
Without question, strenuous exercise is one of the most necessary because neurons are totally intolerant of ischemia.
demanding conditions the cardiovascular system faces. Ulti- Also, the brain is unable to store essential nutrients despite
mately, the major factor determining how long muscles can being the most metabolically active organ in the body.
contract vigorously is the ability of the cardiovascular system Cerebral blood flow is regulated by one of the body’s most
to deliver adequate oxygen and nutrients and remove waste precise autoregulatory systems and is tailored to local neuronal
products. need. For example, when you make a fist with your right hand,
When muscles become active, blood flow increases (hyper- the neurons in the left cerebral motor cortex controlling that
emia) in direct proportion to their greater metabolic activity, movement receive more blood than the adjoining neurons.
a phenomenon called active hyperemia (Figure 19.17). This Brain tissue is exceptionally sensitive to declining pH, and
form of autoregulation occurs almost entirely in response to the increased blood carbon dioxide levels (resulting in acidic con-
decreased oxygen concentration and accumulated metabolic ditions in brain tissue) cause marked vasodilation. Low blood
factors that result from the “revved-up” metabolism of working levels of oxygen are a much less potent stimulus for autoregula-
muscles. tion. However, very high carbon dioxide levels abolish autoreg-
However, systemic adjustments mediated by the vasomo- ulatory mechanisms and severely depress brain activity.
tor center must also occur to ensure that blood delivery to the Besides metabolic controls, the brain also has a myogenic
muscles is both faster and more abundant. During exercise, mechanism that protects it from possibly damaging changes
sympathetic nervous system activity increases. Norepinephrine in blood pressure. When MAP declines, cerebral vessels dilate
released from sympathetic nerve endings causes vasoconstric- to ensure adequate brain perfusion. When MAP rises, cerebral
tion of arterioles throughout the body. This temporarily diverts vessels constrict, protecting the small, more fragile vessels far-
blood away from most regions not essential for exercise, ensur- ther along the pathway from excessive pressure. Under certain
ing that there is sufficient blood pressure to supply the muscles. circumstances, such as brain ischemia caused by rising intra-
In skeletal muscles, the sympathetic nervous system and cranial pressure (as with a brain tumor), the brain (via the med-
local metabolic controls have opposing effects on arteriolar ullary cardiovascular centers) regulates its own blood flow by
diameter. During exercise, local controls override sympathetic triggering a rise in systemic blood pressure.
vasoconstriction. Consequently, blood flow to skeletal muscles However, when systemic pressure changes are extreme, the
brain becomes vulnerable. Fainting, or syncope (sin9cuh-pe;
“cutting short”), occurs when MAP falls below 60 mm Hg.
Cerebral edema is the usual result of pressures over 160 mm
Exercising
skeletal
Hg, which dramatically increase brain capillary permeability.
muscle
The Skin
19
17
Blood flow through the skin:
O2, CO2, H1, and ●● Supplies nutrients to cells
other metabolic factors
in extracellular fluid ●● Helps regulate body temperature
●● Provides a blood reservoir
Autoregulation serves the first function in response to the need
Vasodilation of
arterioles
for oxygen, but the other two require neural intervention. The
(overrides primary function of the cutaneous circulation is to help main-
extrinsic tain body temperature.
sympathetic
input)
Below the skin surface are extensive venous plexuses (net-
works of intertwining vessels). The blood flow through these
plexuses can change from 50 ml/min to as much as 2500 ml/
Initial stimulus
min, depending on body temperature. This capability reflects
Muscle blood
flow (active
Physiological neural adjustments of blood flow through arterioles and through
response
hyperemia)
Result
*Epinephrine acting at beta (b) adrenergic receptors and acetylcholine acting at
cholinergic receptors were once thought to contribute to arteriolar dilation dur-
Figure 19.17 Active hyperemia. Blood flow in exercising ing exercise. However, these appear to have little physiological importance in
skeletal muscle is largely controlled by metabolic autoregulation. controlling human skeletal muscle blood flow.
M19_MARI0853_11_GE_C19.indd 760 03/07/18 2:06 PM

unique coiled arteriovenous anastomoses. These tiny arteriovenous shunts are located
mainly in the fingertips, palms of the hands, toes, soles of the feet, ears, nose, and lips.
Richly supplied with sympathetic nerve endings (unlike the shunts of some other capil-
lary beds), they are controlled by reflexes initiated by temperature receptors or signals
from higher CNS centers. The arterioles, in addition, respond to metabolic autoregula-
tory stimuli.
When the skin is exposed to heat, or body temperature rises for other reasons (such
as vigorous exercise), the hypothalamic “thermostat” signals for reduced vasomotor
stimulation of the skin vessels. As a result, warm blood flushes into the capillary beds
and heat radiates from the skin surface. (This is why Figure 19.15 shows an increase in
blood flow to the skin during exercise.)
When the ambient temperature is cold and body temperature drops, superficial skin
vessels strongly constrict. As a result, blood almost entirely bypasses the capillaries
associated with the arteriovenous anastomoses, diverting the warm blood to the deeper,
more vital organs. Paradoxically, the skin may stay quite rosy because some blood
gets “trapped” in the superficial capillary loops as the shunts swing into operation. The
trapped blood remains red because the chilled skin cells take up less O2.
The Lungs
Blood flow through the pulmonary circuit to and from the lungs is unusual in many
ways. The pathway is relatively short, and pulmonary arteries and arterioles are struc-
turally like veins and venules. That is, they have thin walls and large lumens. Because
resistance to blood flow is low in the pulmonary arterial system, less pressure is needed
to propel blood through those vessels. Consequently, arterial pressure in the pulmonary
circulation is much lower than in the systemic circulation (24/10 versus 120/80 mm Hg).
In the pulmonary circulation, the autoregulatory mechanism is the opposite of what
is seen in most tissues: Low pulmonary oxygen levels cause local vasoconstriction,
and high levels promote vasodilation. While this may seem odd, it is perfectly consis
tent with the gas exchange role of this circulation. When the air sacs of the lungs are
flooded with oxygen-rich air, the pulmonary capillaries become flushed with blood and
ready to receive the oxygen load. In contrast, if the air sacs are collapsed or blocked
with mucus, the oxygen content in those areas is low, and blood largely bypasses those
nonfunctional areas.
The Heart
Aortic pressure and the pumping activity of the ventricles influence the movement of
blood through the smaller vessels of the coronary circulation. When the ventricles con-
tract and compress the coronary blood vessels, blood flow through the myocardium 19
17
stops. As the heart relaxes, the high aortic pressure forces blood through the coronary
circulation.
Under normal circumstances, the myoglobin in cardiac cells stores sufficient oxygen
to satisfy the cells’ oxygen needs during systole. However, an abnormally rapid heart-
beat seriously reduces the ability of the myocardium to receive adequate oxygen and
nutrients during diastole.
Under resting conditions, blood flow through the heart is about 250 ml/min and is
controlled by a myogenic mechanism. Consequently, blood flow remains fairly con-
stant despite wide variations (50 to 140 mm Hg) in coronary perfusion pressure. During
strenuous exercise, the coronary vessels dilate in response to local accumulation of
vasodilators (particularly adenosine), and blood flow may increase three to four times
(see Figure 19.15). Additionally, any event that decreases the oxygen content of the
blood releases vasodilators that adjust the O2 supply to the O2 demand.
This enhanced blood flow during increased heart activity is important because under
resting conditions, cardiac cells use as much as 65% of the oxygen carried to them in
blood. (Most other tissue cells use about 25% of the delivered oxygen.) Consequently,
increasing the blood flow is the only way to provide more oxygen to a vigorously work-
ing heart.
M19_MARI0853_11_GE_C19.indd 761 03/07/18 2:06 PM

Check Your Understanding Velocity in this case is inversely related to cross-sectional area.
17. Suppose you are in a bicycle race. What happens to the
The same thing happens with blood flow inside our blood vessels.
smooth muscle in the arterioles supplying your leg muscles? As shown in Figure 19.18, the speed or velocity of blood
What is the key mechanism in this case? flow changes as blood travels through the systemic circula-
18. If many arterioles in your body dilated at once, you would tion. It is fastest in the aorta and other large arteries (the river),
expect MAP to plummet. What prevents MAP from decreasing slowest in the capillaries (whose large total cross-sectional area
during your bicycle race? makes them analogous to the lake), and then picks up speed
For answers, see Answers Appendix. again in the veins (the river again).
Just as in our analogy of the river and lake, blood flows fastest
where the total cross-sectional area is least. As the arterial sys-
tem branches, the total cross-sectional area of the vascular bed
19.10 Slow blood flow through increases, and the velocity of blood flow declines proportionately.
capillaries promotes diffusion of Even though the individual branches have smaller lumens, their
combined cross-sectional areas and thus the volume of blood they
nutrients and gases, and bulk flow can hold are much greater than that of the aorta.
of fluids For example, the cross-sectional area of the aorta is 2.5 cm2,
Learning Outcome but the combined cross-sectional area of all the capillaries is
4500 cm2. This difference results in fast blood flow in the aorta
NN Outline factors involved in capillary exchange and bulk (40–50 cm/s) and slow blood flow in the capillaries (about
flow, and explain the significance of each.
0.03 cm/s). Slow capillary flow is beneficial because it allows
adequate time for exchanges between the blood and tissue cells.
Velocity of Blood Flow
Have you ever watched a swift river emptying into a large lake?
Vasomotion
The water’s speed decreases as it enters the lake until its flow Blood flow through capillaries is not only slow, it is also inter-
becomes almost imperceptible. This is because the total cross- mittent. The intermittent flow of blood through a capillary bed
sectional area of the lake is much larger than that of the river. is due to vasomotion, the on/off constriction/dilation of arteri-
oles, mostly in response to local chemical conditions (intrinsic
control). Precapillary sphincters also respond to the same local
autoregulatory signals that affect arteriolar diameter.
Relative cross- Capillary Exchange of Respiratory Gases

sectional area of
different vessels and Nutrients
of the vascular bed
Oxygen, carbon dioxide, most nutrients, and metabolic wastes
pass between the blood and interstitial fluid by diffusion. Recall
5000 that in diffusion, net movement always occurs along a concen-
Total area 4000 tration gradient—each substance moving from an area of its
(cm2) of the 3000
higher concentration to an area of its lower concentration. As
19
17 vascular 2000
bed a result, oxygen and nutrients pass from the blood, where their
1000
concentration is fairly high, through the interstitial fluid to the
0
tissue cells. Carbon dioxide and metabolic wastes leave the cells,
50
where their content is higher, and diffuse into the capillary blood.
40
Velocity of There are four different routes across capillaries for different
30
blood flow types of molecules, as Figure 19.19 shows. 1 Lipid-soluble
(cm/s) 20
10
molecules, such as respiratory gases, diffuse through the lipid
0
bilayer of the endothelial cell plasma membranes. Recall that
the plasma membrane is a major barrier to diffusion of solutes
r ta
ies
Ca ioles
ies
les
ins
ae
that are not lipid soluble ( p. 102). Small water-soluble sol-

av
Ao
nu
Ve
ter
lar
ter
ec
Ve
pil
Ar
utes, such as amino acids and sugars, pass through 2 fluid-

Ar
na
filled intercellular capillary clefts or 3 fenestrations. 4 Some

Ve
larger molecules, such as proteins, are actively transported in

Figure 19.18 Blood flow velocity and total cross-sectional
area of vessels. Various blood vessels of the systemic circulation pinocytotic vesicles.
differ in their total cross-sectional area (e.g., the cross section of all As we mentioned earlier, capillaries differ in their “leaki-
systemic capillaries combined versus the cross section of all systemic ness,” or permeability. Liver capillaries, for instance, are sinu-
arteries combined), which affects the velocity of blood flow through soids that allow even proteins to pass freely, whereas brain
them. capillaries are impermeable to most substances.
M19_MARI0853_11_GE_C19.indd 762 03/07/18 2:06 PM

direction and amount of flow across capillary walls reflect the
Pinocytotic balance between two dynamic and opposing forces—hydrostatic
vesicles and colloid osmotic pressures.
Red blood
cell in lumen Hydrostatic Pressures
Endothelial Hydrostatic pressure (HP) is the force exerted by a fluid press-
cell ing against a wall. In capillaries, hydrostatic pressure is the same
as capillary blood pressure—the pressure exerted by blood on
Fenestration capillary walls. Capillary hydrostatic pressure (HPc) tends to
(pore) force fluids through capillary walls (a process called filtration),
Endothelial cell nucleus
leaving behind cells and most proteins. Blood pressure drops as
Basement membrane blood flows along a capillary bed, so HPc is higher at the arterial
Tight
Intercellular
end of the bed (35 mm Hg) than at the venous end (17 mm Hg).
junction In theory, blood pressure—which forces fluid out of the
cleft
capillaries—is opposed by the interstitial fluid hydrostatic
pressure (HPif) acting outside the capillaries and pushing fluid
in. However, there is usually very little fluid in the interstitial
Lumen
space because the lymphatic vessels constantly withdraw it.
Pinocytotic HPif may vary from slightly negative to slightly positive, but
vesicles traditionally it is assumed to be zero.
Endothelial
Colloid Osmotic Pressures
Intercellular fenestration Colloid osmotic pressure (OP), the force opposing hydrostatic
cleft (pore)
pressure, is created by large nondiffusible molecules, such as
4 Transport
via vesicles plasma proteins, that are unable to cross the capillary wall
(large ( p. 676). Such molecules draw water toward themselves. In
substances) other words, they encourage osmosis. This is because water
moves to make the solute more dilute. A quick and dirty way
3 Movement
to remember this is “hydrostatic pressure pushes and osmotic
through
fenestrations pressure sucks.”
(water-soluble The abundant plasma proteins in capillary blood (primarily
substances) albumin molecules) develop a capillary colloid osmotic pres-
2 Movement sure (OPc), also called oncotic pressure, of approximately 26 mm
1 Diffusion through intercellular
clefts (water-soluble Hg. The interstitial fluid colloid osmotic pressure (OPif) is
through plasma
membrane substances) substantially lower—from 0.1 to 5 mm Hg—because interstitial
(lipid-soluble fluid contains few proteins. Unlike HP, OP does not vary signifi-
substances) cantly from one end of the capillary bed to the other.
19
17
Figure 19.19 Capillary transport mechanisms. The four pos- Hydrostatic-Osmotic Pressure Interactions
sible pathways or routes of transport across the endothelial cell wall We are now ready to calculate the net filtration pressure
of a fenestrated capillary.
(NFP), which considers all the forces acting at the capillary
bed. As you work your way through the right-hand page of
Focus Figure 19.1, notice that while net filtration is occurring
Fluid Movements: Bulk Flow at the arteriolar end of the capillary, a negative value for NFP at
the venous end of the capillary indicates that fluid is moving into
While nutrient and gas exchanges are occurring across the cap-
the capillary bed (a process called reabsorption). As a result,
illary walls by diffusion, bulk fluid flows are also going on.
net fluid flow is out of the circulation at the arterial ends of
Fluid is forced out of the capillaries through the clefts at the
capillary beds and into the circulation at the venous ends.
arterial end of the bed, but most of it returns to the bloodstream
However, more fluid enters the tissue spaces than returns to
at the venous end. Though relatively unimportant to capil-
the blood, resulting in a net loss of fluid from the circulation
lary exchange of nutrients and wastes, bulk flow is extremely
of about 1.5 ml/min. Lymphatic vessels pick up this fluid and
important in determining the relative fluid volumes in the
any leaked proteins and return it to the vascular system, which
bloodstream and the interstitial space. (Approximately 20 L of
accounts for the relatively low levels of both fluid and proteins
fluid filter out of the capillaries each day before being returned
in the interstitial space. Were this not so, this “insignificant” fluid
to the blood—almost seven times the total plasma volume!)
loss would empty your blood vessels of plasma in about 24 hours!
As we describe next and show in Focus on Bulk Flow across
Capillary Walls (Focus Figure 19.1 on pp. 764–765), the
(Text continues on p. 766.)
M19_MARI0853_11_GE_C19.indd 763 03/07/18 2:06 PM

FOCUS FIGURE 19.1 Bulk Flow across Capillary Walls
Bulk fluid flow across capillary walls causes continuous mixing of fluid
between the plasma and the interstitial fluid compartments, and maintains
the interstitial environment.
Arteriole
The big picture
Each day, 20 L of fluid filters from capillaries at their
arteriolar end and flows through the interstitial space.
Most (17 L) is reabsorbed at the venous end.
Fluid moves
through the
interstitial space.
For all capillary

beds, 20 L of fluid
is filtered out per
Recall from Chapter 3 ( pp. 103–104) that two kinds day—almost 7
of pressure drive fluid movement: times the total
plasma volume!
Hydrostatic pressure (HP) Osmotic pressure (OP)
• Due to fluid pressing against a • Due to nondiffusible solutes
boundary (e.g., capillary wall) that cannot cross the boundary
• HP “pushes” fluid across the • OP “pulls” fluid across the
boundary boundary
• In blood vessels, is due to • In blood vessels, is due to
blood pressure plasma proteins
Piston
Solute
molecules
(proteins)
Boundary Boundary
“Pushes” “Sucks”
17 L of fluid per
day is reabsorbed
into the capillaries
at the venous end.
About 3 L per
day of fluid
(and any leaked
proteins) are
removed by the
lymphatic
Venule system (see
Chapter 20). Lymphatic
capillary
764
M19_MARI0853_11_GE_C19.indd 764 05/07/18 5:22 PM

How do the pressures drive fluid flow across a capillary?
Net filtration occurs at the arteriolar end of a capillary.

Capillary lumen Boundary Interstitial fluid
(capillary wall)
Hydrostatic pressure in capillary (HPc )

HPc 5 35 mm Hg
“pushes” fluid out of capillary.
Osmotic pressure in capillary (OPc )

OPc 5 26 mm Hg
“pulls” fluid into capillary.
Let’s use what we know about pressures
to determine the net filtration pressure
(NFP) at any point. (NFP is the pressure
Hydrostatic pressure driving fluid out of the capillary.) To do
HPif 5 0 mm Hg (HPif ) in interstitial fluid this we calculate the outward pressures
“pushes” fluid into (HPc and OPif ) minus the inward
capillary. pressures (HPif and OPc ). So,
OPif 5 1 mm Hg Osmotic pressure (OPif ) NFP 5 (HPc 1 OPif ) 2 (HPif 1 OPc )

in interstitial fluid “pulls”
fluid out of capillary. 5 (35 1 1) 2 (0 1 26)
5 10 mm Hg (net outward pressure)
As a result, fluid moves from the

NFP 5 10 mm Hg capillary into the interstitial space.
Net reabsorption occurs at the venous end of a capillary.

Capillary lumen Boundary Interstitial fluid
(capillary wall)
Hydrostatic pressure in capillary
“pushes” fluid out of capillary. HPc 5 17 mm Hg
The pressure has dropped because
of resistance encountered along
the capillaries.
Osmotic pressure in capillary

OPc 5 26 mm Hg
“pulls” fluid into capillary.
Again, we calculate the NFP:

HPif 5 0 mm Hg Hydrostatic pressure in
interstitial fluid “pushes”
fluid into capillary. NFP 5 (HPc 1 OPif ) 2 (HPif 1 OPc )
5 (17 1 1) 2 (0 1 26)
OPif 5 1 mm Hg Osmotic pressure in 5 28 mm Hg (net inward pressure)
interstitial fluid “pulls”
fluid out of capillary. Notice that the NFP at the venous end is
a negative number. This means that
reabsorption, not filtration, is occurring
and so fluid moves from the interstitial
NFP5 28 mm Hg space into the capillary.
When bulk flow goes wrong, edema can result (see Homeostatic Imbalance 19.2, p. 766).
765
M19_MARI0853_11_GE_C19.indd 765 05/07/18 5:22 PM

HOMEOSTATIC
CLINICAL
IMBALANCE 19.2
Edema is an abnormal increase in the amount of interstitial
fluid. You will encounter it frequently in the clinic because it
occurs in diverse clinical scenarios. However, it will be easy
for you to discern the underlying cause of edema in any given
situation if you think of it in terms of the pressures that drive
bulk flow. Either an increase in outward pressure (driving fluid
out of the capillaries) or a decrease in inward pressure could be
the cause.
●● An increase in capillary hydrostatic pressure accelerates
fluid loss from the blood. This could result from incompetent
venous valves, localized blood vessel blockage, congestive
heart failure, or high blood volume. It could also result from
the enlarged uterus of a pregnant woman pressing on veins Figure 19.20 Pitting edema. Applying pressure with a thumb
that return blood to the heart. Whatever the cause, the abnor- leaves an indentation that remains for some time.
mally high capillary hydrostatic pressure intensifies filtration.
●● Increased interstitial fluid osmotic pressure can result from are compensated for by renal mechanisms that maintain blood
an inflammatory response. Inflammation increases capil- volume and pressure. However, rapid onset of edema such as
lary permeability, allowing plasma proteins to leak into the that in anaphylaxis may have serious effects on the efficiency
interstitial fluid. Together, the more porous capillaries and of the circulation due to a decrease in blood volume and blood
the increased osmolality of the interstitial fluid draw large pressure.
amounts of fluid out of the capillaries, accounting for the
localized swelling seen in inflammation. In an anaphylac- Check Your Understanding
tic response (see p. 845), edema results from the massive
19. DRAW At a given point in a capillary, suppose that capillary
release of the inflammatory chemical histamine.
hydrostatic pressure is 32 mm Hg, interstitial fluid hydrostatic
●● Decreased capillary colloid osmotic pressure hinders fluid pressure is 1 mm Hg, capillary colloid osmotic pressure is
return to the blood. Since plasma proteins are largely respon- 25 mm Hg, and interstitial fluid osmotic pressure is 2 mm
sible for OPc, hypoproteinemia (hi0po-pro0te-ĭ-ne9me-ah), a Hg. Draw a line representing the capillary wall and label
condition of unusually low levels of plasma proteins, results the compartments on either side as “capillary lumen” and
in tissue edema. Fluids are forced out of the capillary beds “interstitial fluid.” For each of the four pressures, draw an
at the arteriolar ends by blood pressure as usual, but fail to arrow across the capillary wall pointing in the correct direction,
return to the blood at the venous ends. As a result, the inter- and label it with the appropriate name and value. Calculate
the net filtration pressure. Would you expect to find this point
stitial spaces become congested with fluid. Hypoprotein-
at the venous or arterial end of the capillary?
emia may result from liver disease, protein malnutrition, or
20. PREDICT Suppose OPif rises dramatically—say because of a
glomerulonephritis (in which plasma proteins pass through
severe bacterial infection in the surrounding tissue. (a) Predict
19
17 “leaky” renal filtration membranes and are lost in urine).
how fluid flow will change in this situation. (b) Now calculate
●● Theoretically, a decrease in interstitial fluid hydrostatic the NFP at the venous end of the capillary in Focus Figure 19.1
pressure should also be a potential cause of edema. How- if OPif increases to 10 mm Hg. (c) In which direction does fluid
ever, this does not occur because HPif is too low to decrease flow at the venous end of the capillary now—in or out?
to any extent. 21. MAKE CONNECTIONS Your patient in right heart failure is
●● A fourth cause of edema is decreased drainage of intersti- experiencing peripheral edema. Which of the four pressures
that drive bulk fluid flow at capillaries has been changed as a
tial fluid through lymphatic vessels that have been blocked
result of the heart failure and in which direction?
(e.g., by parasitic worms; see elephantiasis in the Chapter 20
Related Clinical Terms on p. 809) or surgically removed (for For answers, see Answers Appendix.
example, during cancer surgery).
Edema can occur anywhere in the body but is most easily PART 3
visible in the skin. Excess interstitial fluid in the subcutaneous
tissues generally causes pitting edema (Figure 19.20). Gravity CIRCULATORY PATHWAYS:
determines where edematous fluid accumulates, so involvement BLOOD VESSELS OF THE BODY
of the legs and feet is common.
Edema can impair tissue function because excess fluid in Learning Outcomes
the interstitial space increases the distance nutrients and oxy- NN Trace the pathway of blood through the pulmonary
gen must diffuse between the blood and the cells. Usually circuit, and state the importance of this special
edema develops slowly, and so the fluid losses from the blood circulation.
M19_MARI0853_11_GE_C19.indd 766 03/07/18 2:06 PM

NN Describe the general functions of the systemic circuit. tive organs enters a special subcirculation, the hepatic portal
The term vascular system is often used to describe the body’s system, and perfuses through the liver before it reenters the
complex network of blood vessels. However, as we saw in general systemic circulation (see Table 19.12).
Chapter 18, the heart is actually a double pump that serves two
distinct circulations, each with its own set of arteries, capil-
laries, and veins. The pulmonary circulation is the short loop The vessels of the systemic
19.11
that runs from the heart to the lungs and back to the heart. The
systemic circulation routes blood through a long loop to all
circulation transport blood to all
parts of the body before returning it to the heart. Table 19.3 on body tissues
pp. 768–769 shows both circuits schematically. Learning Outcomes
The heart pumps all of its blood into a single systemic
NN Name and give the location of the major arteries and
artery—the aorta. In contrast, blood returning to the heart is
veins in the systemic circulation.
delivered largely by two terminal systemic veins, the superior NN Describe the structure and special function of the hepatic
and inferior venae cavae. The single exception to this is the portal system.
blood draining from the myocardium of the heart, which is col-
lected by the cardiac veins and reenters the right atrium via the The principal arteries and veins of the systemic circulation are
coronary sinus. described in Tables 19.4 through 19.13. (The fetal circulation
In addition to these differences between arteries and veins is described later in the Developmental Aspects section of this
connecting to the heart, there are three important differences chapter and in Chapter 28.)
between systemic arteries and veins: Notice that by convention, oxygen-rich blood is shown red,
while blood that is relatively oxygen-poor is depicted blue,
●● Arteries run deep while veins are both deep and superficial.
regardless of vessel type. The schematic flowcharts (pipe dia-
Deep veins parallel the course of the systemic arteries and
grams) that accompany each table show the vessels that would
both are protected by body tissues along most of their course.
be closer to the viewer in brighter, more intense colors than
With a few exceptions, these veins are named identically to
vessels deeper or farther from the viewer. For example, darker
their companion arteries. Superficial veins run just beneath the
blue veins would be closer to the viewer than lighter blue veins
skin and are readily seen, especially in the limbs, face, and
in the body region shown.
neck. Because there are no superficial arteries, the names of
As you examine the tables that follow and locate the various
the superficial veins do not correspond to the names of any of
systemic arteries and veins in the illustrations, be aware of cues
the arteries.
that make your memorization task easier. Also notice that:
●● Venous pathways are more interconnected. Unlike the
fairly distinct arterial pathways, venous pathways tend to ●● In many cases, the name of a vessel reflects the body region
have numerous interconnections, and many veins are rep- traversed (axillary, brachial, femoral, etc.), the organ served
resented by not one but two similarly named vessels. As a (renal, hepatic, gonadal), or the bone followed (vertebral,
result, venous pathways are more difficult to follow. radial, tibial).
●● The brain and digestive systems have unique venous
●● Arteries and veins tend to run side by side and, in many
drainage systems. Most body regions have a similar pat- places, they also run with nerves.
tern for their arterial supply and venous drainage. However, ●● The systemic vessels do not always match on the right and left 19
17
the venous drainage patterns in at least two important body sides of the body. Thus, while almost all vessels in the head
areas are unique. First, venous blood draining from the brain and limbs are bilaterally symmetrical, some of the large, deep
enters large dural venous sinuses ( pp. 496–497) rather vessels of the trunk region are asymmetrical or unpaired.
than typical veins. Second, blood draining from the diges-
M19_MARI0853_11_GE_C19.indd 767 03/07/18 2:06 PM

Table 19.3 Pulmonary and Systemic Circulations
Pulmonary Circulation
The pulmonary circulation (Figure 19.21a) functions only to Pulmonary Pulmonary
capillaries R. pulmonary L. pulmonary capillaries
bring blood into close contact with the alveoli (air sacs) of the of the of the
artery artery
lungs so that gases can be exchanged. It does not directly serve R. lung L. lung
the metabolic needs of body tissues.
Oxygen-poor, dark red blood enters the pulmonary circula-
tion as it is pumped from the right ventricle into the large pul-
monary trunk (Figure 19.21b), which runs diagonally upward To
for about 8 cm and then divides abruptly to form the right and Pulmonary
systemic
trunk
left pulmonary arteries. In the lungs, the pulmonary arteries circulation
subdivide into the lobar arteries (lo9bar) (three in the right lung
and two in the left lung), each of which serves one lung lobe. R. pulmonary veins
The lobar arteries accompany the main bronchi into the lungs
and then branch profusely, forming first arterioles and then the
dense networks of pulmonary capillaries that surround and
cling to the delicate air sacs. It is here that oxygen moves from From
the alveolar air to the blood and carbon dioxide moves from the systemic RA LA
blood to the alveolar air. As gases are exchanged and the oxy- circulation
gen content of the blood rises, the blood becomes bright red. L. pulmonary
The pulmonary capillary beds drain into venules, which join veins
RV LV
to form the two pulmonary veins exiting from each lung. The
four pulmonary veins complete the circuit by unloading their
precious cargo into the left atrium of the heart. (a) Schematic flowchart.
Left pulmonary
artery Air-filled
alveolus
Aortic arch of lung
Pulmonary trunk
Right pulmonary O2
artery
Three lobar arteries CO2

to right lung Pulmonary
capillary
19
17 Gas exchange
Two lobar arteries

to left lung
Pulmonary
veins
Pulmonary
Right veins
atrium
Left atrium
Right
ventricle
Left ventricle
(b) Illustration. The pulmonary arterial system is shown in blue to indicate that the blood it carries is oxygen-poor.
The pulmonary venous drainage is shown in red to indicate that the blood it transports is oxygen-rich.
Figure 19.21 Pulmonary circulation. (RA 5 right atrium, RV 5 right ventricle, LA 5 left atrium, LV 5 left ventricle)
M19_MARI0853_11_GE_C19.indd 768 03/07/18 2:06 PM

Table 19.3 (continued)
Note that any vessel with the term pulmonary or lobar in its
name is part of the pulmonary circulation. All others are part of
the systemic circulation. Common
Capillary beds of carotid arteries
Pulmonary arteries carry oxygen-poor, carbon dioxide–rich to head and
head and
blood, and pulmonary veins carry oxygen-rich blood.* This is upper limbs subclavian
opposite to the systemic circulation, where arteries carry oxy- arteries to
upper limbs
gen-rich blood and veins carry carbon dioxide–rich, relatively Superior
oxygen-poor blood. vena cava Aortic
arch
Systemic Circulation
The systemic circulation provides the functional blood supply to
all body tissues; that is, it delivers oxygen, nutrients, and other
needed substances while carrying away carbon dioxide and Aorta
other metabolic wastes. Freshly oxygenated blood* returning
from the pulmonary circuit is pumped out of the left ventricle
into the aorta (Figure 19.22).
From the aorta, blood can take various routes, because
essentially all systemic arteries branch from this single great
vessel. The aorta arches upward from the heart and then curves
and runs downward along the body midline to its terminus in RA LA
the pelvis, where it splits to form the two large arteries serv-
ing the lower extremities. The branches of the aorta continue
to subdivide to produce the arterioles and, finally, the capil-
RV LV
laries that spread throughout the organs. Venous blood drain-
ing from organs inferior to the diaphragm ultimately enters Azygos Thoracic
system aorta
the inferior vena cava.† Except for some coronary and thoracic
venous drainage (which enters the azygos system of veins), the Venous Arterial
superior vena cava drains body regions above the diaphragm. drainage blood
The venae cavae empty the carbon dioxide–laden blood into the
right atrium of the heart.
Two important points concerning the two major circulations:
Inferior
●● Blood passes from systemic veins to systemic arteries vena Capillary beds of
cava mediastinal structures
only after first moving through the pulmonary circuit
and thorax walls
(Figure 19.21a).
●● Although the entire cardiac output of the right ventricle Diaphragm
19
17
passes through the pulmonary circulation, only a small frac- Abdominal
tion of the output of the left ventricle flows through any aorta
single organ (Figure 19.22).
The systemic circulation can be viewed as multiple circulatory
channels functioning in parallel to distribute blood to all body Capillary beds of
Inferior digestive viscera,
organs. vena spleen, pancreas,
cava kidneys
*
By convention, oxygen-rich blood is shown red and oxygen-poor blood is
shown blue.
†
Venous blood from the digestive viscera passes through the hepatic portal circu-
lation (liver and associated veins) before entering the inferior vena cava.
Capillary beds of
gonads, pelvis, and
lower limbs
Figure 19.22 Schematic flowchart showing an overview of

the systemic circulation. The pulmonary circulation is shown
in gray for comparison. (RA 5 right atrium, RV 5 right
ventricle, LA 5 left atrium, LV 5 left ventricle)
M19_MARI0853_11_GE_C19.indd 769 03/07/18 2:06 PM

Table 19.4 The Aorta and Major Arteries of the Systemic Circulation
Figure 19.23a diagrams the distribution of the aorta and major arteries, which supply the myocardium. The aortic arch, deep
arteries of the systemic circulation in flowchart form, and Fig- to the sternum, begins and ends at the sternal angle (T4 level).
ure 19.23b illustrates them. See Tables 19.5 through 19.8 for Its three major branches (R to L) are: (1) the brachiocephalic
fine points about the vessels arising from the aorta. trunk (bra9ke-o-sĕ-fal0ik; “armhead”), which passes superiorly
The aorta (a-or9tah) is the largest artery in the body. In adults, under the right sternoclavicular joint and branches into the right
the aorta is approximately the size of a garden hose where it common carotid artery (kah-rot9id) and the right subclavian
issues from the left ventricle of the heart. Its internal diameter artery, (2) the left common carotid artery, and (3) the left sub-
is 2.5 cm, and its wall is about 2 mm thick. It decreases in size clavian artery. These three vessels provide the arterial supply of
slightly as it runs to its terminus. The aortic valve guards the the head, neck, upper limbs, and part of the thorax wall.
base of the aorta and prevents backflow of blood during diastole. The descending aorta runs along the anterior spine. Called
Opposite each aortic valve cusp is an aortic sinus, which contains the thoracic aorta from T5 to T12, it sends off numerous small
baroreceptors important in reflex regulation of blood pressure. arteries to the thorax wall and viscera before piercing the
Different portions of the aorta are named according to shape diaphragm. As it enters the abdominal cavity, it becomes the
or location. The first portion, the ascending aorta, runs posteri- abdominal aorta. This portion supplies the abdominal walls
orly and to the right of the pulmonary trunk. It persists for only and viscera and ends at the L4 level, where it splits into the
about 5 cm before curving to the left as the aortic arch. The only right and left common iliac arteries, which supply the pelvis
branches of the ascending aorta are the right and left coronary and lower limbs.
R. external R. internal L. external L. internal Figure 19.23 Major arteries of the

carotid artery carotid artery carotid artery carotid artery systemic circulation.
R. common carotid L. common carotid

R. vertebral L. vertebral
• Right side of head and neck • Left side of head and neck
R. axillary R. subclavian Brachiocephalic L. subclavian L. axillary

• Neck and • Head, neck, and • Neck and L.
R. upper limb R. upper limb Aortic arch upper limb
Arteries of Arteries of
R. upper L. upper
Ascending aorta limb
limb
L. and R. coronary Thoracic aorta
arteries
L. ventricle of heart
Visceral branches Parietal branches
Mediastinal Esophageal Bronchial Pericardial Posterior intercostals Superior phrenics

19
17 • Posterior • Esophagus • Lungs and • Pericardium • Intercostal muscles, spinal • Posterior and superior
media- bronchi cord, vertebrae, pleurae, skin diaphragm
stinum
Diaphragm
Abdominal aorta
Visceral branches Parietal branches
Gonadal Suprarenal Superior Celiac trunk Inferior phrenics Lumbars Median sacral
• Testes or • Adrenal and inferior • Liver • Inferior diaphragm • Posterior • Sacrum
ovaries glands mesenterics • Gallbladder abdominal • Coccyx
and • Small • Spleen wall
Renal intestine • Stomach
• Kidneys • Colon • Esophagus
• Duodenum
R. common iliac L. common iliac

• Pelvis and R. lower limb • Pelvis and L. lower limb
Arteries of R. lower limb Arteries of L. lower limb
(a) Schematic flowchart
M19_MARI0853_11_GE_C19.indd 770 03/07/18 2:06 PM

Arteries of the head and trunk

Internal carotid artery
External carotid artery
Common carotid arteries
Vertebral artery Arteries that supply the upper limb
Subclavian artery Subclavian artery
Brachiocephalic trunk
Aortic arch Axillary artery
Ascending aorta
Coronary artery
Celiac trunk Brachial artery

Abdominal aorta
Superior mesenteric artery
Renal artery
Radial artery
Gonadal artery
Ulnar artery
Inferior mesenteric artery
Common iliac artery
Internal iliac artery
Deep palmar arch
Superficial palmar arch
Digital arteries
Arteries that supply the lower limb

External iliac artery 19
17
Femoral artery
Popliteal artery
Anterior tibial artery
Posterior tibial artery
Arcuate artery
(b) Illustration, anterior view
Figure 19.23 (continued)
M19_MARI0853_11_GE_C19.indd 771 03/07/18 2:06 PM

Table 19.5 Arteries of the Head and Neck

Four paired arteries supply the head and neck. These are the Each common carotid divides into two major branches (the
common carotid arteries, plus three branches from each subcla- internal and external carotid arteries). At the division point,
vian artery: the vertebral arteries, the thyrocervical trunks, and each internal carotid artery has a slight dilation, the carotid
the costocervical trunks (Figure 19.24b). Of these, the common sinus, that contains baroreceptors that assist in reflex blood
carotid arteries have the broadest distribution (Figure 19.24a). pressure control. The carotid bodies, chemoreceptors involved
in controlling respiratory rate, are located close by. Pressing
on the neck in the area of the carotid sinuses can cause uncon-
R. anterior L. anterior
cerebral artery cerebral artery
sciousness (carot 5 stupor) because the pressure created mim-
ics high blood pressure, eliciting vasodilation, which interferes
R. middle with blood delivery to the brain.
cerebral
artery Description and Distribution
Anterior
communicating
artery Common carotid arteries. The origins of these two arteries
Cerebral arterial circle
differ: The right common carotid artery arises from the bra-
chiocephalic trunk; the left is the second branch of the aortic
R. and L.
posterior arch. The common carotid arteries ascend through the lateral
Ophthalmic communicating neck, and at the superior border of the larynx (the level of the
artery arteries “Adam’s apple”), each divides into its two major branches, the
external and internal carotid arteries.
R. posterior Basilar
cerebral artery
The external carotid arteries supply most tissues of the
Superficial head except for the brain and orbit. As each artery runs superi-
artery
temporal
artery orly, it sends branches to the thyroid gland and larynx (superior
thyroid artery), the tongue (lingual artery), the skin and mus-
cles of the anterior face (facial artery), and the posterior scalp
Maxillary R. and L. (occipital artery). Each external carotid artery terminates by
artery vertebral
arteries splitting into a superficial temporal artery, which supplies the
parotid salivary gland and most of the scalp, and a maxillary
Occipital artery, which supplies the upper and lower jaws and chewing
artery
muscles, the teeth, and the nasal cavity. A clinically important
R. and L. branch of the maxillary artery is the middle meningeal artery
Facial internal
artery carotid (not illustrated). It enters the skull through the foramen spino-
arteries sum and supplies the inner surface of the parietal bone, squa-
Lingual mous part of the temporal bone, and the underlying dura mater.
artery The larger internal carotid arteries supply the orbits and
R. and L. more than 80% of the cerebrum. They assume a deep course
19
17 Superior external and enter the skull through the carotid canals of the temporal
thyroid carotid
arteries
bones (Figure 7.6a and b, p. 238). Once inside the cranium,
artery
each artery gives off one main branch, the ophthalmic artery,
and then divides into the anterior and middle cerebral arteries.
R. and L. The ophthalmic arteries (of-thal9mik) supply the eyes, orbits,
common forehead, and nose. Each anterior cerebral artery supplies the
carotid
arteries medial surface of the frontal and parietal lobes of the cerebral
R. and L.
hemisphere on its side and also anastomoses with its partner on
subclavian the opposite side via a short arterial shunt called the anterior
arteries communicating artery (Figure 19.24c). The middle cerebral
arteries run in the lateral sulci of their respective cerebral hem-
ispheres and supply the lateral parts of the temporal, parietal,
Aortic arch and frontal lobes.
Vertebral arteries. The vertebral arteries spring from the
subclavian arteries at the root of the neck and ascend through
(a) Schematic flowchart foramina in the transverse processes of the cervical vertebrae
to enter the skull through the foramen magnum. En route, they
Figure 19.24 Arteries of the head, neck, and brain.
M19_MARI0853_11_GE_C19.indd 772 03/07/18 2:06 PM

send branches to the vertebrae and cervical spinal

cord and to some deep structures of the neck. Within
Ophthalmic artery the cranium, the right and left vertebral arteries join
to form the basilar artery (bas9ĭ-lar), which ascends
along the anterior aspect of the brain stem, giving
Branches of off branches to the cerebellum, pons, and inner ear
the external
Basilar artery carotid artery
(Figure 19.24b and c). At the pons-midbrain border,
Vertebral • Superficial
the basilar artery divides into a pair of posterior cere
artery temporal artery bral arteries, which supply the occipital lobes and
Internal • Maxillary artery the inferior parts of the temporal lobes.
carotid artery • Occipital artery Arterial shunts called posterior communicating
External • Facial artery arteries connect the posterior cerebral arteries to the
carotid artery
• Lingual artery
middle cerebral arteries anteriorly. The two posterior
Common arteries and a single anterior communicating artery
• Superior thyroid
carotid artery
artery complete the formation of an arterial anastomosis
Thyrocervical called the cerebral arterial circle (circle of Willis).
trunk This structure encircles the pituitary gland and optic
Larynx
Costocervical
Thyroid gland
chiasma and unites the brain’s anterior and posterior
trunk blood supplies. It also equalizes blood pressure in
(overlying trachea)
Subclavian
Clavicle (cut) the two brain areas and provides alternate routes for
artery
Brachiocephalic blood to reach the brain tissue if a carotid or vertebral
Axillary trunk artery becomes occluded.
artery
Internal thoracic Thyrocervical and costocervical trunks. These short
artery
vessels arise from the subclavian artery just lateral
to the vertebral arteries on each side (Figure 19.24b
(b) Arteries of the head and neck, right aspect and Figure 19.25). The thyrocervical trunk mainly
supplies the thyroid
Anterior gland, portions of the
Cerebral arterial cervical vertebrae and
Frontal lobe circle
(circle of Willis)
spinal cord, and some
Optic chiasma scapular muscles. The
• Anterior costocervical trunk
communicating
Middle artery serves deep neck and
cerebral
artery
superior intercostal
• Anterior muscles.
cerebral artery
19
17
Internal • Posterior
carotid communicating
artery artery
Mammillary • Posterior
body cerebral artery
Temporal Basilar artery

lobe
Vertebral artery
Pons
Occipital lobe
Cerebellum
Posterior
(c) Major arteries serving the brain (inferior view, right side
of cerebellum and part of right temporal lobe removed)
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Table 19.6 Arteries of the Upper Limbs and Thorax
The upper limbs are supplied entirely by arteries arising from an array of vessels that arise either directly from the thoracic
the subclavian arteries (Figure 19.25a). After giving off aorta or from branches of the subclavian arteries. Most visceral
branches to the neck, each subclavian artery courses laterally organs of the thorax receive their functional blood supply from
between the clavicle and first rib to enter the axilla, where its small branches issuing from the thoracic aorta. Because these
name changes to axillary artery. The thorax wall is supplied by vessels are so small and tend to vary in number (except for the
bronchial arteries), Figure 19.25a and b does not illustrate them,
but several are listed at the end of this table.
R. common L. common carotid Description and Distribution

carotid artery
R. vertebral artery Arteries of the Upper Limb
artery
Thyrocervical trunk L. vertebral artery Axillary artery. As it runs through the axilla accompa-
L. subclavian nied by cords of the brachial plexus, each axillary artery
Suprascapular artery
artery gives off branches to the axilla, chest wall, and shoulder
R. subclavian artery girdle. These branches include the thoracoacromial
Axillary artery artery (tho0rah-ko-ah-kro9me-al), which supplies the
Thoracoacromial deltoid muscle and pectoral region; the lateral thoracic
artery artery, which serves the lateral chest wall and breast; the
Thoracoacromial subscapular artery to the scapula, dorsal thorax wall,
artery Aortic arch Costocervical and part of the latissimus dorsi muscle; and the ante-
(pectoral trunk
branch)
rior and posterior circumflex humeral arteries, which
wrap around the humeral neck and help supply the shoul-
Anterior Brachiocephalic der joint and the deltoid muscle. As the axillary artery
and posterior trunk emerges from the axilla, it becomes the brachial artery.
circumflex
humeral Internal Brachial artery. The brachial artery runs down the medial
arteries thoracic aspect of the humerus and supplies the anterior flexor mus-
artery
cles of the arm. One major branch, the deep artery of the
Brachial
artery Anterior arm, serves the posterior triceps brachii muscle. As it
intercostal nears the elbow, the brachial artery gives off several small
arteries
branches that contribute to an anastomosis serving the
Deep Lateral elbow joint and that connect to the arteries of the forearm.
artery thoracic
artery Thoracic Posterior As the brachial artery crosses the anterior midline aspect
of arm aorta intercostal
Subscapular
of the elbow, it provides an easily palpated pulse point
arteries
artery (brachial pulse) (see Figure 19.8). Immediately beyond the
elbow, the brachial artery splits to form the radial and ulnar
Anastomosis
arteries, which more or less follow the course of similarly
19
17 Common named bones down the anterior forearm.
interosseous artery
Radial artery. The radial artery runs from the median
line of the cubital fossa to the styloid process of the
radius. It supplies the lateral muscles of the forearm, the
Radial Ulnar artery
artery
wrist, and the thumb and index finger. Proximal to the
carpal bones, the radial artery provides a convenient site
for taking the radial pulse.
Ulnar artery. The ulnar artery supplies the medial aspect
of the forearm, fingers 3–5, and the medial aspect of the
index finger. Proximally, the ulnar artery gives off a short
Deep
palmar branch, the common interosseous artery (in0ter-os9e-us),
arch Metacarpal arteries which runs between the radius and ulna to serve the deep
Superficial palmar arch flexors and extensors of the forearm.
Palmar arches. In the palm, branches of the radial and
Digital arteries ulnar arteries anastomose to form the superficial and deep
(a) Schematic flowchart palmar arches. The metacarpal arteries and digital
arteries that supply the fingers arise from these palmar
Figure 19.25 Arteries of the right upper limb and thorax. arches.
M19_MARI0853_11_GE_C19.indd 774 03/07/18 2:06 PM

Common carotid arteries

Vertebral artery
Thyrocervical trunk Right subclavian artery
Costocervical trunk
Suprascapular artery Left subclavian artery
Thoracoacromial artery
Axillary artery
Subscapular artery
Posterior circumflex Posterior intercostal
humeral artery arteries
Anterior circumflex
humeral artery Anterior intercostal
artery
Brachial artery Internal thoracic artery
Lateral thoracic artery

Deep artery of arm
Descending aorta
Common interosseous
artery
Radial artery the intercostal spaces anteriorly. The internal thoracic artery
also sends superficial branches to the skin and mammary
Ulnar artery glands and terminates in twiglike branches to the anterior
abdominal wall and diaphragm.
Posterior intercostal arteries. The superior two pairs of pos-
terior intercostal arteries are derived from the costocervical
trunk. The next nine pairs issue from the thoracic aorta and
course around the rib cage to anastomose anteriorly with the
anterior intercostal arteries. Inferior to the 12th rib, a pair of
Deep palmar arch subcostal arteries emerges from the thoracic aorta (not illus- 19
17
trated). The posterior intercostal arteries supply the posterior
Superficial palmar arch intercostal spaces, deep muscles of the back, vertebrae, and spi-
nal cord. Together, the posterior and anterior intercostal arteries
Digital arteries
supply the intercostal muscles.
Superior phrenic arteries. One or more paired superior
phrenic arteries serve the posterior superior aspect of the dia-
phragm surface.
(b) Illustration, anterior view Arteries of the Thoracic Viscera
Pericardial arteries. Several tiny branches supply the poste-
Figure 19.25 (continued) rior pericardium.
Bronchial arteries. Two left and one right bronchial arteries
Arteries of the Thorax Wall supply systemic (oxygen-rich) blood to the lungs, bronchi, and
Internal thoracic arteries. The internal thoracic arteries (for- pleurae.
merly called the internal mammary arteries) arise from the Esophageal arteries. Four to five esophageal arteries supply
subclavian arteries and supply blood to most of the anterior the esophagus.
thorax wall. Each of these arteries descends lateral to the ster- Mediastinal arteries. Many small mediastinal arteries serve
num and gives off anterior intercostal arteries, which supply the posterior mediastinum.
M19_MARI0853_11_GE_C19.indd 775 03/07/18 2:06 PM

Table 19.7 Arteries of the Abdomen
The arterial supply to the abdominal organs arises from the mesenteric arteries, and the median sacral artery, all are paired
abdominal aorta (Figure 19.26a). Under resting conditions, vessels. These arteries supply the abdominal wall, diaphragm,
about half of the entire arterial flow moves through these and visceral organs of the abdominopelvic cavity. We discuss
vessels. Except for the celiac trunk, the superior and inferior the branches in the order of their issue.
Diaphragm
Abdominal L. gastric artery
aorta
Inferior R. gastric
phrenic artery
arteries Hepatic
Common
hepatic artery L
artery proper
Celiac
trunk
Gastro-
Splenic duodenal R
artery artery
R. gastroepiploic
artery
Middle L. gastroepiploic artery
suprarenal Intestinal arteries
arteries
Middle colic
artery
Superior
mesenteric
artery R. colic
artery
Renal
arteries
Ileocolic artery
Gonadal
arteries Sigmoidal
19
17 arteries
Inferior
mesenteric
artery
L. colic
artery
Superior rectal
Lumbar artery
arteries
Median sacral artery
Common iliac arteries
(a) Schematic flowchart.
Figure 19.26 Arteries of the abdomen.
M19_MARI0853_11_GE_C19.indd 776 03/07/18 2:06 PM

Description and Distribution ●● Splenic artery. As the splenic artery (splen9ik) passes deep
Inferior phrenic arteries. The inferior phrenics emerge from to the stomach, it sends branches to the pancreas and stom-
the aorta at T12, just inferior to the diaphragm (Figure 19.26c). ach and terminates in branches to the spleen.
They serve the inferior diaphragm surface. ●● Left gastric artery. The left gastric artery (gaster 5 stom-
Celiac trunk. This very large unpaired branch of the abdomi- ach) supplies part of the stomach and the inferior esophagus.
nal aorta divides almost immediately into three branches The right and left gastroepiploic arteries (gas0tro-ep0ĭ-
(Figure 19.26b): plo9ik)—branches of the gastroduodenal and splenic arteries,
●● Common hepatic artery. The common hepatic artery respectively—serve the greater curvature of the stomach. A
(hĕ-pat9ik) gives off branches to the stomach, duodenum, and right gastric artery, which supplies the stomach’s lesser cur-
pancreas. Where the gastroduodenal artery branches off, vature, may arise from the common hepatic artery or from the
the common hepatic becomes the hepatic artery proper, hepatic artery proper.
which splits into right and left branches that serve the liver.
Liver (cut) Diaphragm
Inferior vena cava

Esophagus
Celiac trunk
Left gastric artery
Stomach
Common hepatic artery
Splenic artery
Hepatic artery proper
Left gastroepiploic
Gastroduodenal artery artery
Right gastric artery
Spleen
Gallbladder
Pancreas Right gastroepiploic
(major portion lies artery
posterior to stomach)
Duodenum
Abdominal aorta Superior mesenteric
artery
(b) The celiac trunk and its major branches. The left half of the liver has been removed. 19
17
➤
M19_MARI0853_11_GE_C19.indd 777 03/07/18 2:06 PM

Table 19.7 Arteries of the Abdomen (continued)
Superior mesenteric artery (mes-en-ter9ik). This large, males. The ovarian arteries extend into the pelvis to serve the
unpaired artery arises from the abdominal aorta at the L1 level ovaries and part of the uterine tubes. The much longer testicular
immediately below the celiac trunk (Figure 19.26d). It runs arteries descend through the pelvis and inguinal canals to enter
deep to the pancreas and then enters the mesentery (a drapelike the scrotum, where they serve the testes.
membrane that supports the small intestine), where its numer- Inferior mesenteric artery. This final major branch of the
ous anastomosing branches serve virtually all of the small intes- abdominal aorta is unpaired and arises from the anterior aor-
tine via the intestinal arteries, most of the large intestine—the tic surface at the L3 level. It serves the distal part of the large
appendix, cecum, ascending colon (via the ileocolic and right intestine—from the midpart of the transverse colon to the
colic arteries)— and part of the transverse colon (via the mid- midrectum—via its left colic, sigmoidal, and superior rectal
dle colic artery). branches (Figure 19.26d). Looping anastomoses between the
Suprarenal arteries (soo0prah-re9nal). The middle suprare- superior and inferior mesenteric arteries help ensure that blood
nal arteries flank the origin of the superior mesenteric artery will continue to reach the digestive viscera in cases of trauma
as they emerge from the abdominal aorta (Figure 19.26c). They to one of these abdominal arteries.
supply blood to the adrenal (suprarenal) glands overlying the Lumbar arteries. Four pairs of lumbar arteries arise from the
kidneys. The adrenal glands also receive two sets of branches posterolateral surface of the aorta in the lumbar region (Figure
not illustrated: superior suprarenal branches from the nearby 19.26c). These segmental arteries supply the posterior abdomi-
inferior phrenic arteries, and inferior suprarenal branches from nal wall.
the nearby renal arteries.
Median sacral artery. The unpaired median sacral artery
Renal arteries. The short but wide renal arteries, right and issues from the posterior surface of the abdominal aorta at its
left, issue from the lateral surfaces of the aorta slightly below terminus. This tiny artery supplies the sacrum and coccyx.
the superior mesenteric artery (between L1 and L2). Each serves
Common iliac arteries. At the L4 level, the aorta splits into the
the kidney on its side.
right and left common iliac arteries, which supply blood to the
Gonadal arteries (go-nă9dul). The paired gonadal arteries are lower abdominal wall, pelvic organs, and lower limbs.
called ovarian arteries in females and testicular arteries in
Diaphragm
Hiatus (opening)
for inferior vena cava
Inferior phrenic artery
Hiatus (opening)
for esophagus
Adrenal (suprarenal) Middle suprarenal artery

gland
Celiac trunk
19
17 Renal artery
Kidney
Superior mesenteric artery
Abdominal aorta
Gonadal (testicular
or ovarian) artery
Lumbar arteries
Inferior mesenteric artery

Ureter
Median sacral
artery
Common iliac artery
(c) Major branches of the abdominal aorta.
Figure 19.26 (continued) Arteries of the abdomen.
M19_MARI0853_11_GE_C19.indd 778 03/07/18 2:06 PM

Transverse colon
Celiac trunk
Abdominal aorta
Superior mesenteric
artery
Inferior mesenteric
artery
Branches of the superior
mesenteric artery
• Middle colic artery
Branches of the inferior
• Intestinal arteries mesenteric artery
• Right colic artery • Left colic artery
• Ileocolic artery • Sigmoidal arteries
• Superior rectal artery
Ascending colon
Right common iliac artery

Descending colon
Ileum
Cecum
Sigmoid colon
Appendix
Rectum
(d) Distribution of the superior and inferior mesenteric arteries. The transverse colon has been pulled superiorly.
19
17
22. Which paired artery supplies most of the tissues of the head
except for the brain and orbits?
23. Name the arterial anastomosis at the base of the cerebrum.
24. Name the four unpaired arteries that emerge from the
abdominal aorta.
M19_MARI0853_11_GE_C19.indd 779 03/07/18 2:06 PM

Table 19.8 Arteries of the Pelvis and Lower Limbs
At the level of the sacroiliac joints, the common iliac arteries

Abdominal
aorta divide into two major branches, the internal and external iliac
Superior arteries (Figure 19.27a). The internal iliacs distribute blood
gluteal mainly to the pelvic region. The external iliacs primarily serve
artery the lower limbs but also send branches to the abdominal wall.
Internal
iliac
Inferior
artery
Description and Distribution
gluteal Common
artery iliac Internal iliac arteries. These paired arteries run into the pelvis
artery and distribute blood to the pelvic walls and viscera (bladder
Internal Obturator and rectum, plus the uterus and vagina in the female and the
pudendal artery prostate and ductus deferens in the male). Additionally they
Deep artery
Medial of thigh External serve the gluteal muscles via the superior and inferior gluteal
circumflex iliac arteries, adductor muscles of the medial thigh via the obtura-
femoral artery
artery
tor artery, and external genitalia and perineum via the internal
pudendal artery (not illustrated).
Lateral External iliac arteries. These arteries supply the lower limbs
circumflex
femoral Femoral (Figure 19.27b). As they course through the pelvis, they give off
artery artery branches to the anterior abdominal wall. After passing under the
inguinal ligaments to enter the thigh, they become the femoral
arteries.
Adductor Femoral arteries. As each of these arteries passes down the
hiatus
anteromedial thigh, it gives off several branches to the thigh
muscles. The largest of the deep branches is the deep artery
of the thigh (also called the deep femoral artery), which is the
Arterial main supply to the thigh muscles (hamstrings, quadriceps, and
anastomosis
adductors). Proximal branches of the deep femoral artery, the
lateral and medial circumflex femoral arteries, encircle the
Popliteal neck of the femur. The medial circumflex artery is the major
artery
vessel to the head of the femur. If it is torn in a hip fracture, the
bone tissue of the head of the femur dies. A long descending
Anterior Posterior tibial branch of the lateral circumflex artery supplies the vastus later-
tibial artery alis muscle. Near the knee the femoral artery passes posteriorly
artery
and through a gap in the adductor magnus muscle, the adductor
hiatus, to enter the popliteal fossa, where its name changes to
popliteal artery.
19
17
Fibular Popliteal artery. This posterior vessel contributes to an arte-
(peroneal) rial anastomosis that supplies the knee region and then splits
artery
into the anterior and posterior tibial arteries of the leg.
Anterior tibial artery. The anterior tibial artery runs through
Dorsalis the anterior compartment of the leg, supplying the extensor
pedis Lateral muscles along the way. At the ankle, it becomes the dorsal-
artery plantar
artery
is pedis artery, which supplies the ankle and dorsum of the
foot, and gives off a branch, the arcuate artery, which issues
Lateral
plantar the dorsal metatarsal arteries to the metatarsus of the foot.
Medial
artery plantar The superficial dorsalis pedis ends by penetrating into the sole
Arcuate artery where it forms the medial part of the plantar arch. The dorsal-
artery is pedis artery provides a clinically important pulse point, the
Plantar arch
pedal pulse. If the pedal pulse is easily felt, it is fairly certain
that the blood supply to the leg is good.
Dorsal Plantar
metatarsal metatarsal
arteries arteries
(a) Schematic flowchart Figure 19.27 Arteries of the right pelvis and lower limb.
M19_MARI0853_11_GE_C19.indd 780 03/07/18 2:06 PM

Posterior tibial artery. This large artery courses through the

posteromedial part of the leg and supplies the flexor muscles.
Common iliac artery Proximally, it gives off a large branch, the fibular (peroneal)
Internal iliac artery artery, which supplies the lateral fibularis muscles of the leg.
On the medial side of the foot, the posterior tibial artery divides
Superior gluteal artery
into lateral and medial plantar arteries that serve the plantar
External iliac artery surface of the foot. The lateral plantar artery forms the lateral
end of the plantar arch. Plantar metatarsal arteries and digital
Deep artery of thigh
arteries to the toes arise from the plantar arch.
Lateral circumflex
femoral artery
Medial circumflex
femoral artery
Obturator artery
Femoral artery
Adductor hiatus
Popliteal artery
Popliteal artery

Fibular artery
Fibular artery
Lateral plantar artery 19
17
Dorsalis pedis artery
(from top of foot)
Dorsalis pedis artery
Medial plantar artery
Arcuate artery
Dorsal metatarsal Plantar arch

arteries
(b) Anterior view (c) Posterior view
Check Your Understanding locations: behind the knee, behind the medial malleolus of the
tibia, on the dorsum of the foot.
25. You are assessing the circulation in the leg of a diabetic patient
at the clinic. Name the artery you palpate in each of these three For answers, see Answers Appendix.
M19_MARI0853_11_GE_C19.indd 781 03/07/18 2:06 PM

Table 19.9 The Venae Cavae and the Major Veins of the Systemic Circulation
In our survey of the systemic veins, the major tributaries (branches) heart wall. It is formed by the union of the right and left bra-
of the venae cavae are noted first in Figure 19.28, followed by a chiocephalic veins and empties into the right atrium (Figure
description in Tables 19.10 through 19.13 of the venous pattern of 19.28b). Notice that there are two brachiocephalic veins, but
the various body regions. Because veins run toward the heart, the only one brachiocephalic artery (trunk). Each brachiocephalic
most distal veins are named first and those closest to the heart last. vein is formed by the joining of the internal jugular and sub-
Deep veins generally drain the same areas served by their com- clavian veins on its side. In most of the flowcharts that follow,
panion arteries, so they are not described in detail. only the vessels draining blood from the right side of the body
are shown (except for the azygos circulation of the thorax).
Description and Areas Drained
Inferior vena cava. The widest blood vessel in the body, this
Superior vena cava. This great vein receives systemic blood vein returns blood to the heart from all body regions below the
draining from all areas superior to the diaphragm, except the diaphragm. The abdominal aorta lies directly to its left. The
paired common iliac veins join at L5 to form the distal
R. external R. vertebral Intracranial end of the inferior vena cava. From this point, it courses
Veins of
R. upper
jugular • Cervical spinal dural venous sinuses superiorly along the anterior aspect of the spine, receiv-
• Superficial cord and ing venous blood from the abdominal walls, gonads,
limb
head and neck vertebrae
kidneys, adrenal glands, and liver. Immediately above
the diaphragm, the inferior vena cava ends as it enters
R. internal jugular
• Dural venous the inferior aspect of the right atrium.
sinuses of the brain
R. subclavian
R. axillary • R. head, neck,
and upper
Same as R. brachiocephalic
limb
R. brachiocephalic L. brachiocephalic
• R. side of head and R. upper limb • L. side of head and L. upper limb
Superior vena cava Azygos system

• Drains much of thorax
R. atrium of heart
Diaphragm
Inferior vena cava
L., R., and middle

19
17 hepatic veins
R. suprarenal • Liver
(L. suprarenal drains
into L. renal vein)
• Adrenal glands L. and R. renal veins
• Kidneys
R. gonadal
(L. gonadal drains Lumbar veins
into L. renal vein) (several pairs)
• Testis or ovary • Posterior abdominal
wall
R. common iliac L. common iliac

• Pelvis and R. lower • Pelvis and L. lower
limb limb
Veins of Veins of
R. lower limb L. lower limb
Figure 19.28 Major veins of the systemic circulation.
M19_MARI0853_11_GE_C19.indd 782 03/07/18 2:06 PM

Veins of the head and trunk

Dural venous sinuses
External jugular vein

Vertebral vein Veins that drain
the upper limb
Internal jugular vein
Subclavian vein
Right and left
brachiocephalic veins Axillary vein
Superior vena cava Cephalic vein

Brachial vein
Great cardiac vein Basilic vein
Hepatic veins
Splenic vein
Hepatic portal vein
Renal vein
Median cubital vein
Superior mesenteric vein
Inferior mesenteric vein Ulnar vein
Inferior vena cava Radial vein
Common iliac vein
Internal iliac vein
Digital veins
Veins that drain the lower limb

External iliac vein
Femoral vein 19
17
Great saphenous vein
Popliteal vein
Posterior tibial vein
Anterior tibial vein

Small saphenous vein
Dorsal venous arch

Dorsal metatarsal veins
(b) Illustration, anterior view
M19_MARI0853_11_GE_C19.indd 783 03/07/18 2:06 PM

Table 19.10 Veins of the Head and Neck
Three pairs of veins collect most of the blood draining from the sagittal sinuses are in the falx cerebri, which dips down between
head and neck (Figure 19.29a): the cerebral hemispheres. The inferior sagittal sinus drains into
●● The external jugular veins, which empty into the subclavians the straight sinus posteriorly (Figure 19.29a and c). The supe-
rior sagittal and straight sinuses then empty into the transverse
●● The internal jugular veins, which join with the subclavians sinuses, which run in shallow grooves on the internal surface
●● The vertebral veins, which drain into the brachiocephalic of the occipital bone. These drain into the S-shaped sigmoid
veins sinuses, which become the internal jugular veins as they leave
Although most extracranial veins have the same names as the the skull through the jugular foramen. The cavernous sinuses,
extracranial arteries, their courses and interconnections differ which flank the sphenoid body, receive venous blood from the
substantially. ophthalmic veins of the orbits and the facial veins, which drain
Most veins of the brain drain into the dural venous sinuses, the nose and upper lip area. The internal carotid artery and cra-
an interconnected series of enlarged chambers located between nial nerves III, IV, VI, and part of V all run through the cavern-
the dura mater layers ( p. 496). The superior and inferior ous sinus on their way to the orbit and face.
Description and Area Drained

External jugular veins. The right and left external
Superior sagittal sinus
jugular veins drain superficial scalp and face structures
served by the external carotid arteries. However, their
tributaries anastomose frequently, and some of the
Inferior
sagittal sinus Superficial superficial drainage from these regions enters the inter-
temporal vein nal jugular veins as well. As the external jugular veins
descend through the lateral neck, they pass obliquely
Straight Ophthalmic over the sternocleidomastoid muscles and then empty
sinus vein into the subclavian veins.
Occipital Transverse
vein Vertebral veins. Unlike the vertebral arteries, the ver-
sinus
tebral veins do not serve much of the brain. Instead they
Cavernous drain the cervical vertebrae, the spinal cord, and some
sinus Facial vein
small neck muscles. They run inferiorly through the
transverse foramina of the cervical vertebrae and join
Posterior auricular the brachiocephalic veins at the root of the neck.
vein
Internal jugular veins. The paired internal jugular
Sigmoid sinus Internal jugular vein veins, which receive the bulk of blood draining from
External jugular
the brain, are the largest of the paired veins drain-
Superior thyroid vein
vein ing the head and neck. They arise from the dural
Vertebral vein
venous sinuses, exit the skull via the jugular foramina
19
17 Middle thyroid vein
( Figure 7.6a and b, p. 238), and then descend through
Brachiocephalic veins the neck alongside the internal carotid arteries. As they
move inferiorly, they receive blood from some of the
Subclavian vein
deep veins of the face and neck—branches of the facial
and superficial temporal veins (Figure 19.29b). At the
Superior vena cava base of the neck, each internal jugular vein joins the
(a) Schematic flowchart subclavian vein on its own side to form a brachio
cephalic vein. As already noted, the two brachiocephalic
Figure 19.29 Venous drainage of the head, neck, and brain. veins unite to form the superior vena cava.
M19_MARI0853_11_GE_C19.indd 784 03/07/18 2:06 PM

Ophthalmic vein
Superficial
temporal vein
Facial vein
Occipital vein
Posterior
auricular vein
External
jugular vein
Vertebral vein
Internal
jugular vein
Superior and middle

thyroid veins
Brachiocephalic
vein
Subclavian vein
Superior
vena cava
(b) Veins of the head and neck, right superficial aspect
Superior sagittal sinus
Falx cerebri
Inferior sagittal sinus
Straight sinus
Cavernous sinus
19
17
Transverse sinuses
Sigmoid sinus
Jugular foramen
Right internal jugular vein
(c) Dural venous sinuses of the brain
Check Your Understanding 27. Which veins drain the dural venous sinuses and where do
these veins terminate?
26. In what important way does the area drained by the vertebral
veins differ from the area served by the vertebral arteries? For answers, see Answers Appendix.
M19_MARI0853_11_GE_C19.indd 785 03/07/18 2:06 PM

Table 19.11 Veins of the Upper Limbs and Thorax
Subclavian Internal The deep veins of the upper limbs follow the paths of their
vein jugular vein companion arteries and have the same names (Figure 19.30a).
External Brachiocephalic
jugular vein veins However, except for the largest, most are paired veins that flank
Axillary
vein their artery. The superficial veins of the upper limbs are larger
than the deep veins and are easily seen just beneath the skin.
The median cubital vein, crossing the anterior aspect of the
elbow, is commonly used to obtain blood samples or administer
Superior
vena cava
intravenous medications.
Blood draining from the mammary glands and the first two
to three intercostal spaces enters the brachiocephalic veins.
However, the vast majority of thoracic tissues and the thorax
wall are drained by a complex network of veins called the azy-
gos system (a-zi9gos). The branching nature of the azygos sys-
tem provides a collateral circulation for draining the abdominal
Accessory wall and other areas served by the inferior vena cava, and there
hemiazygos are numerous anastomoses between the azygos system and the
vein
inferior vena cava.
Median
cubital
vein Description and Areas Drained
Deep Veins of the Upper Limbs
The most distal deep veins of the upper limb are the radial and
Brachial Azygos Hemiazygos ulnar veins. The deep and superficial venous palmar arches
vein vein vein
of the hand empty into the radial and ulnar veins of the fore-
Right and left posterior arm, which then unite to form the brachial vein of the arm. As
intercostal veins the brachial vein enters the axilla, it becomes the axillary vein,
which becomes the subclavian vein at the level of the first rib.
Superficial Veins of the Upper Limbs
The superficial venous system begins with the dorsal venous net-
work (not illustrated), a plexus of superficial veins in the dorsum
of the hand. In the distal forearm, this plexus drains into two
major superficial veins—the cephalic and basilic veins—which
anastomose frequently as they course upward (Figure 19.30b).
The cephalic vein bends around the radius as it travels superi-
Cephalic Median Basilic orly and then continues up the lateral superficial aspect of the
vein antebrachial vein
19
17 vein
arm to the shoulder, where it runs in the groove between the del-
toid and pectoralis muscles to join the axillary vein. The basilic
vein courses along the posteromedial aspect of the forearm,
Radial Ulnar
vein vein crosses the elbow, and then joins the brachial vein in the axilla,
forming the axillary vein. At the anterior aspect of the elbow,
Deep venous the median cubital vein connects the basilic and cephalic veins.
palmar arch The median antebrachial vein lies between the radial and ulnar
Metacarpal veins veins in the forearm and terminates (variably) at the elbow by
entering either the basilic or the cephalic vein.
Superficial venous
palmar arch The Azygos System
Digital veins The azygos system consists of the following vessels, which
flank the vertebral column laterally.
Azygos vein. Located against the right side of the vertebral
Figure 19.30 Veins of the thorax and right upper limb. For column, the azygos vein (azygos 5 unpaired) originates in the
clarity, the abundant branching and anastomoses of the superficial abdomen, from the right ascending lumbar vein that drains
veins are not shown. most of the right abdominal cavity wall and from the right pos-
terior intercostal veins (except the first) that drain the chest
muscles. At the T4 level, it arches over the great vessels that run
to the right lung and empties into the superior vena cava.
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Hemiazygos vein (hĕ0me-a-zi9gos; “half the azygos”). This Accessory hemiazygos vein. The accessory hemiazygos com-
vessel ascends on the left side of the vertebral column. Its ori- pletes the venous drainage of the left (middle) thorax and can
gin, from the left ascending lumbar vein and the lower (9th– be thought of as a superior continuation of the hemiazygos vein.
11th) posterior intercostal veins, mirrors that of the inferior It receives blood from the 4th–8th posterior intercostal veins
portion of the azygos vein on the right. About midthorax, the and then crosses to the right to empty into the azygos vein. Like
hemiazygos vein passes in front of the vertebral column and the azygos, it receives oxygen-poor systemic blood from the
joins the azygos vein. bronchi of the lungs (bronchial veins).
Brachiocephalic veins Internal jugular vein

External jugular vein
Right subclavian vein
Left subclavian vein

Axillary vein
Superior vena cava

Brachial vein
Azygos vein
Cephalic vein
Accessory hemiazygos vein
Basilic vein
Hemiazygos vein
Posterior intercostals
Inferior vena cava
Median cubital vein Ascending lumbar vein
Median
antebrachial
19
17
vein
Basilic vein
Cephalic vein
Ulnar vein
Radial vein
Deep venous
palmar arch
Superficial venous
palmar arch
Digital veins
(b) Anterior view
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Table 19.12 Veins of the Abdomen
The inferior vena cava returns blood from the abdominopelvic together—is called a portal system and always serves very spe-
viscera and abdominal walls to the heart (Figure 19.31a). Most cific needs. The hepatic portal system carries nutrient-rich
of its venous tributaries have names that correspond to the arter- blood (which may also contain toxins and microorganisms)
ies serving the abdominal organs. from the digestive organs to the liver, where it can be “treated”
Veins draining the digestive viscera empty into a common before it reaches the rest of the body. As the blood percolates
vessel, the hepatic portal vein, which transports this venous slowly through the liver sinusoid capillaries, hepatocytes pro-
blood into the liver before it is allowed to enter the major cess nutrients and toxins, and phagocytic cells rid the blood of
systemic circulation via the hepatic veins (Figure 19.31c). bacteria and other foreign matter.
Such a venous system—veins connecting two capillary beds
Description and Areas Drained
The veins of the abdomen are listed in inferior to supe-
Inferior rior order.
vena cava
Lumbar veins. Several pairs of lumbar veins drain the
Inferior phrenic veins posterior abdominal wall. They empty both directly into
Cystic vein Hepatic veins the inferior vena cava and into the ascending lumbar
veins of the azygos system of the thorax.
Gonadal (testicular or ovarian) veins. The right
gonadal vein drains the ovary or testis on the right side
of the body and empties into the inferior vena cava. The
left gonadal vein drains into the left renal vein superiorly.
Hepatic Hepatic portal vein
portal Renal veins. The right and left renal veins drain the
system kidneys.
Superior mesenteric vein
Splenic vein Suprarenal veins. The right suprarenal vein drains the
adrenal gland on the right and empties into the inferior
vena cava. The left suprarenal vein drains into the left
Suprarenal Inferior renal vein.
veins mesenteric
vein
Hepatic portal system. Like all portal systems, the
Renal veins
hepatic portal system is a series of vessels in which two
Gonadal veins separate capillary beds lie between the arterial supply
and the final venous drainage. In this case, the first capil-
lary beds are in the stomach and intestines and drain into
tributaries of the hepatic portal vein, which brings them
Lumbar veins to the second capillary bed in the liver. The short hepatic
portal vein begins at the L2 level. Numerous tributaries
19
17 from the stomach and pancreas contribute to the hepatic
portal system (Figure 19.31c), but the major vessels are:
●● Superior mesenteric vein: Drains the entire small
intestine, part of the large intestine (ascending and
R. ascending transverse regions), and stomach.
lumbar vein L. ascending
lumbar vein ●● Splenic vein: Collects blood from the spleen, parts of
the stomach and pancreas, and then joins the superior
mesenteric vein to form the hepatic portal vein.
●● Inferior mesenteric vein: Drains the distal portions
Common iliac veins of the large intestine and rectum and joins the splenic
External iliac vein vein just before that vessel unites with the superior
mesenteric vein to form the hepatic portal vein.
Internal iliac veins Hepatic veins. The right, left, and middle hepatic veins
(a) Schematic flowchart.
carry venous blood from the liver to the inferior vena cava.
Cystic veins. The cystic veins drain the gallbladder
Figure 19.31 Veins of the abdomen. and join the portal veins in the liver.
Inferior phrenic veins. The inferior phrenic veins
drain the inferior surface of the diaphragm.
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Inferior phrenic vein

Hepatic veins
Inferior vena cava

Left suprarenal vein
Right suprarenal vein
Renal veins
Left ascending
Right gonadal vein lumbar vein
Lumbar veins
Left gonadal vein
Common iliac vein
External iliac vein Internal iliac vein
(b) Tributaries of the inferior vena cava. Venous drainage of abdominal organs not drained by the hepatic portal vein.
Hepatic veins
Gastric veins
Liver
Spleen
Inferior vena cava

Hepatic portal vein
Splenic vein 19
17
Right
gastroepiploic vein
Inferior
mesenteric vein
Superior
mesenteric vein
Small intestine
Large intestine
Rectum
(c) The hepatic portal circulation.
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Table 19.13 Veins of the Pelvis and Lower Limbs
As in the upper limbs, most deep veins of the lower limbs have Superficial veins. The great and small saphenous veins
the same names as the arteries they accompany and many are (sah-fe9nus) issue from the dorsal venous arch of the foot
double. Poorly supported by surrounding tissues, the two super- (Figure 19.32b and c). These veins anastomose frequently with
ficial saphenous veins (great and small) are common sites of each other and with the deep veins along their course. The great
varicosities. The great saphenous (saphenous 5 obvious) vein saphenous vein is the longest vein in the body. It travels supe-
is frequently excised and used as a coronary bypass vessel. riorly along the medial aspect of the leg to the thigh, where it
empties into the femoral vein just distal to the inguinal liga-
Description and Areas Drained ment. The small saphenous vein runs along the lateral aspect of
Deep veins. After being formed by the union of the medial the foot and then through the deep fascia of the calf muscles,
and lateral plantar veins, the posterior tibial vein ascends which it drains. At the knee, it empties into the popliteal vein.
deep in the calf muscle and receives the fibular (peroneal)
vein (Figure 19.32). The anterior tibial vein, which is the
superior continuation of the dorsalis pedis vein of the foot,
unites at the knee with the posterior tibial vein to form the
popliteal vein, which crosses the back of the knee. As the pop-
Common iliac vein
liteal vein emerges from the knee, it becomes the femoral vein,
which drains the deep structures of the thigh. The femoral vein Internal iliac vein
becomes the external iliac vein as it enters the pelvis. In the External iliac vein
pelvis, the external iliac vein unites with the internal iliac vein
to form the common iliac vein. The distribution of the internal
Inguinal ligament
iliac veins parallels that of the internal iliac arteries.
Femoral vein
Inferior Great saphenous
Common vena cava vein (superficial)
iliac vein
Internal
iliac vein
External iliac vein
Great
saphenous
Femoral Femoral vein
Great Popliteal
vein vein
saphenous vein Popliteal
vein vein
Small
saphenous Anterior
19
17 Popliteal tibial vein
vein
Small vein Small Fibular
saphenous saphenous Fibular vein
vein vein vein
Small
Anterior Anterior saphenous
Fibular tibial vein
tibial (peroneal) vein
vein vein (superficial)
Fibular Dorsalis
(peroneal) Posterior pedis vein Posterior
vein tibial tibial
vein Dorsal vein
venous
Plantar Plantar
Dorsalis arch
Dorsal veins veins
venous pedis
vein Deep Dorsal
arch Deep
plantar arch metatarsal plantar arch
Dorsal veins
metatarsal Digital
veins Digital veins
veins
Anterior Posterior
(a) Schematic flowchart of the anterior and posterior veins (b) Anterior view (c) Posterior view
Figure 19.32 Veins of the right lower limb.
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Check Your Understanding umbilical vein and arteries, large vessels that circulate blood
28. Below is a schematic drawing of the hepatic portal system
between the fetal circulation and the placenta where gas and
showing two capillary beds separated by a vein. State the nutrient exchanges occur with the mother’s blood (see Chapter
locations of capillary beds a and c. What type of capillaries are 28). Once the fetal circulatory pattern is laid down, few vascular
found in bed c? Name the major veins labeled b, d, and e. changes occur until birth, when the umbilical vessels and shunts
are occluded (blocked).
Aorta Hepatic portal system
Unlike congenital heart diseases, congenital vascular prob-
Nutrients and Nutrients lems are rare, and blood vessels are remarkably trouble-free
toxins absorbed and toxins during youth. Vessels form as needed to support body growth
leave
e and wound healing, and to rebuild vessels lost each month dur-
ing a woman’s menstrual cycle. As we age, signs of vascular
b d disease begin to appear. In some, the venous valves weaken,
a c
and purple, snakelike varicose veins appear. In others, more
insidious signs of inefficient circulation appear: tingling fingers
First capillary bed Second capillary bed
and toes and cramping muscles.
29. Name the leg veins that often become varicosed. Although the degenerative process of atherosclerosis begins
30. MAKE CONNECTIONS You learned about another portal system in youth, its consequences are rarely apparent until middle to
in Chapter 16. Name that portal system. old age, when it may precipitate a myocardial infarction (heart
attack) or stroke. Until puberty, the blood vessels of boys and
girls look alike, but from puberty to about age 45, women have
strikingly less atherosclerosis than men because of the protec-
Developmental Aspects of Blood tive effects of estrogens. Estrogens reduce resistance to blood
flow and increase the production of HDL (“good” lipoprotein),
Vessels which reduces the risk of atherosclerosis.
The endothelial lining of blood vessels is formed by mesodermal Between the ages of 45 and 65, when estrogen production
cells, which collect in little masses called blood islands through- wanes in women, this “gap” between the sexes closes, and males
out the microscopic embryo. These blood islands form fragile and females above age 65 are equally at risk for cardiovascular
sprouting extensions that reach toward one another and toward disease. You might expect that giving postmenopausal women
the forming heart to lay down the rudimentary vascular tubes. supplementary estrogens would maintain this protective effect.
Meanwhile, adjacent mesenchymal cells, stimulated by platelet- Surprisingly, clinical trials have shown that this is not the case.
derived growth factor, surround the endothelial tubes, forming Blood pressure changes with age. In a newborn baby, arterial
the stabilizing muscular and fibrous coats of the vessel walls. pressure is about 90/55. Blood pressure rises steadily during
How do blood vessels “know” where to grow? Many blood childhood to finally reach the adult value (120/80). After age
vessels simply follow the same guidance cues that nerves follow, 40, the incidence of hypertension increases dramatically, as do
which is why forming vessels often snuggle closely to nerves. associated illnesses such as heart attacks, strokes, vascular dis-
Whether a vessel becomes an artery or a vein depends upon the ease, and renal failure.
local concentration of a differentiation factor called vascular At least some vascular disease is a product of our modern
endothelial growth factor. As noted in Chapter 18, the heart technological culture. “Blessed” with high-protein and lipid- 19
17
pumps blood through the rudimentary vascular system by the rich diets, empty-calorie snacks, energy-saving devices, and
fourth week of development. high-stress jobs, many of us are struck down prematurely. Life-
In addition to the fetal shunts that bypass the nonfunctional style modifications—a healthy diet, regular aerobic exercise,
lungs (the foramen ovale and ductus arteriosus), other vascular and eliminating cigarette smoking—can help prevent cardio-
modifications are found in the fetus. A special vessel, the ductus vascular disease.
venosus, largely bypasses the liver. Also important are the
R E L AT E D C L I N I C A L T E R M S
Aneurysm (an9u-rizm; aneurysm 5 a widening) A balloonlike Angiogram (an9je-o-gram0; angio 5 a vessel; gram 5 writing)
outpocketing of an artery wall that places the artery at risk for Diagnostic technique involving the infusion of a radiopaque
rupture; most often reflects gradual weakening of the artery substance into the circulation for X-ray examination of specific
by chronic hypertension or atherosclerosis. The most common blood vessels ( p. 48). The major technique for diagnosing
sites of aneurysms are the abdominal aorta and arteries coronary artery occlusion and risk of a heart attack.
feeding the brain and kidneys. Deep venous thrombosis Clot formation in a deep vein. An ever-
present danger is that the clot may detach and form a life-
threatening pulmonary embolus.
M19_MARI0853_11_GE_C19.indd 791 03/07/18 2:06 PM

A CLOSER LOOK CLINICAL
Atherosclerosis? Get Out the Cardiovascular Drāno®
When pipes get clogged, it is usually immune to its ravages? A large number of
because we’ve dumped something down the interacting risk factors, such as increasing
drain that shouldn’t be there, or something age, being male, family history, high blood
is growing inside (tree roots, for example), cholesterol, hypertension, cigarette smoking,
trapping the normal sludge coming through lack of exercise, diabetes, obesity, stress,
(see top photo). In atherosclerosis, small and intake of trans fats are involved.
patchy thickenings called atheromas form A growing body of evidence links
that make the walls of our arteries thicker systemic inflammation with the formation
and stiffer, resulting in hypertension. and subsequent rupture of atherosclerotic
Atheromas can intrude into the vessel plaques. C-reactive protein is a marker of
lumen, making it easy for arterial spasms or systemic inflammation that is measured to
a blood clot to close the vessel completely. predict the likelihood of future heart attacks
and strokes.
Onset and Stages
Atherosclerosis indirectly causes half of the Prevention and Treatment
deaths in the Western world. How does this Some risk factors are under our control. We
scourge of blood vessels come about? The can avoid smoking, lose weight, exercise
development of a full-blown atheroma is regularly to increase blood levels of high-
believed to occur in several stages. density lipoprotein (HDL, the “good”
1. The endothelium is injured. lipoprotein that removes cholesterol from
The initial event is damage to the vessel walls and carries it to the liver), and eat
endothelium caused by turbulent blood a healthy diet low in saturated and trans fats.
flow, bloodborne chemicals (such as But for many of us, these measures are
cholesterol), hypertension, components not enough. Cholesterol-lowering, anti-
of cigarette smoke, or viral or bacterial inflammatory statins (and even the humble
infections. Damaged endothelial cells aspirin) can help decrease the risk of heart
become dysfunctional, and this sets the attacks and strokes.
stage for atherosclerosis. Plaques that partially block arteries are
Top A pipe clogged by accumulated
2. Lipids accumulate and oxidize in the treated in much the same way we would
deposits. Bottom Atherosclerotic plaques
tunica intima. Injured endothelial cells set treat a blocked sewer pipe—dig it up and
nearly close a human artery.
up a local inflammatory response, attracting replace it or call a plumber to drill through
white blood cells. Endothelial cells then the obstruction. In coronary bypass surgery,
begin to transport and modify low-density complicated plaque, it is unstable and veins or small arteries removed from other
lipoproteins (LDLs) that deliver cholesterol to prone to rupture. areas of the body are implanted in the
tissue cells from the blood. The accumulated heart to restore its circulation. In balloon
LDL oxidizes in the inflammatory Consequences angioplasty, a balloon is passed through the
environment. This not only damages Plaques stiffen artery walls, causing vessels to the obstruction and then inflated
neighboring cells, but also attracts more hypertension. The increased pressure stresses to compress the fatty mass against the
macrophages. Some of these macrophages the plaques, making them even more unstable. vessel wall.
become so engorged with LDLs that they Plaques also constrict the vessel and cause the Angioplasty temporarily clears the path,
are transformed into lipid-laden foam cells. arterial walls to fray and ulcerate, conditions but restenoses (new blockages) often occur.
3. Smooth muscle cells proliferate and a that encourage thrombus (clot) formation. Stents, short metal-mesh tubes, are sometimes
fibrous cap forms. Smooth muscle cells Two other factors also promote thrombus used to help hold the vessel open.
from the tunica media deposit collagen formation: (1) Injured endothelial cells When an atheroma ruptures and
and elastic fibers. The thickened intima, make less anti-thrombotic chemicals. (2) induces clot formation, thrombolytic (clot-
called a fibrous or atherosclerotic Lipoprotein (a), an altered form of LDL found dissolving) agents can help. A genetically
plaque, has a core of dead and dying in some individuals, inhibits fibrinolysis. engineered form of the naturally occurring
foam cells. At first the vessel walls Plaque formation increases the risk of heart tissue plasminogen activator (tPA) is injected
accommodate the growing plaque by attacks and strokes, and is responsible for the directly into the blocked vessel. tPA restores
expanding outward, but eventually these pain (angina) that occurs when heart muscle blood flow quickly and puts an early end to
fatty mounds begin to protrude into is ischemic. Atherosclerosis, particularly of the many heart attacks and strokes in progress.
the vessel lumen, producing full-blown aorta, can also cause aneurysms (ballooning Of course, it’s best to prevent
atherosclerosis (see bottom photo). of the arterial wall) that may burst. atherosclerosis from progressing in the first
4. The plaque becomes unstable. As the place by changing our lifestyles. But how
plaque enlarges, the cells at its center Risk Factors many of us will give up our burgers, butter,
die, calcium is deposited, and collagen Why are some of us so troubled by and sedentary lifestyles for the sake of
fiber production declines. Now called a atherosclerosis while others are seemingly preventing atherosclerosis?
792
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SYSTEM CONNEC TIONS
Homeostatic Interrelationships between the

Cardiovascular System and Other Body Systems
Nervous System Chapters 11–15
●● The cardiovascular system delivers oxygen and nutrients; carries
away wastes
●● The ANS regulates cardiac rate and force; sympathetic division
maintains blood pressure and controls blood flow to skin for
thermoregulation
Endocrine System Chapter 16

away wastes; blood serves as a transport vehicle for hormones
●● Various hormones influence blood pressure (epinephrine, ANP,
angiotensin II, thyroxine, ADH); estrogens maintain vascular
health in premenopausal women
Lymphatic System/Immunity Chapters 20–21

●● The cardiovascular system delivers oxygen and nutrients to
lymphatic organs, which house immune cells; provides transport
medium for lymphocytes and antibodies; carries away wastes
●● The lymphatic system picks up leaked fluid and plasma proteins
and returns them to the cardiovascular system; immune cells
protect cardiovascular organs from specific pathogens
Respiratory System Chapter 22

away wastes
●● The respiratory system carries out gas exchange: loads oxygen
and unloads carbon dioxide from the blood; respiratory
“pump” aids venous return
Digestive System Chapter 23

away wastes
Integumentary System Chapter 5 ●● The digestive system provides nutrients to the blood including
●● The cardiovascular system delivers oxygen and nutrients; carries iron and B vitamins essential for RBC (and hemoglobin)
away wastes formation
●● The skin vasculature is an important blood reservoir and Urinary System Chapters 25–26
provides a site for heat loss from the body 19
17
Skeletal System Chapters 6–8 away wastes; blood pressure drives filtration in the kidneys
●● The urinary system helps regulate blood volume and pressure by
away wastes altering urine volume and releasing renin
●● Bones are the sites of hematopoiesis; protect cardiovascular Reproductive System Chapter 27
organs by enclosure; and provide a calcium depot ●● The cardiovascular system delivers oxygen and nutrients; carries
Muscular System Chapters 9–10 away wastes
●● Estrogens maintain vascular and bone health in women
away wastes
●● The muscular “pump” aids venous return; aerobic exercise
enhances cardiovascular efficiency and helps prevent
atherosclerosis
793
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Diuretic (diure 5 urinate) A chemical that promotes urine Phlebotomy (flĕ-bot9o-me; tomy 5 cut) A venous incision or puncture
formation, thus reducing blood volume. Diuretic drugs are made for the purpose of withdrawing blood or bloodletting.
frequently prescribed to manage hypertension. Sclerotherapy Procedure for removing varicose or spider veins.
Phlebitis (flĕ-bi9tis; phleb 5 vein; itis 5 inflammation) Tiny needles are used to inject scarring agents into the abnormal
Inflammation of a vein accompanied by painful throbbing and vein. The vein scars, closes down, and is absorbed by the body.
redness of the skin over the inflamed vessel. It is most often Superficial thrombophlebitis Inflammation and clot formation in
caused by bacterial infection or local physical trauma. superficial veins, usually in the leg.
C H A P T E R S U M M A RY
PART 1 19.5 Anastomoses are special interconnections between
BLOOD VESSEL STRUCTURE AND FUNCTION blood vessels (p. 746)
1. Blood is transported throughout the body via a continuous 1. The joining together of blood vessels to provide alternate
system of blood vessels. Arteries transport blood away from channels in the same organ is called an anastomosis. Vascular
the heart; veins carry blood back to the heart. Capillaries carry anastomoses form between arteries, between veins, and
blood to tissue cells and are exchange sites. between arterioles and venules.
19.1 Most blood vessel walls have three layers (p. 741) PART 2
1. All blood vessels except capillaries have three layers: tunica
PHYSIOLOGY OF CIRCULATION
intima, tunica media, and tunica externa. Capillary walls are
composed of the tunica intima only. 19.6 Blood flows from high to low pressure against
resistance (pp. 746–747)
19.2 Arteries are pressure reservoirs, distributing vessels,
or resistance vessels (p. 742) 1. Blood flow is the amount of blood flowing through a vessel, an
organ, or the entire circulation in a given period of time. Blood
1. Elastic (conducting) arteries are the large arteries close to the
pressure (BP) is the force per unit area exerted on a vessel wall
heart that expand during systole, acting as pressure reservoirs,
by the contained blood. Resistance is opposition to blood flow;
and then recoil during diastole to keep blood moving. Muscular
blood viscosity and blood vessel length and diameter contribute
(distributing) arteries carry blood to specific organs; they are
to resistance.
less stretchy and more active in vasoconstriction. Arterioles
2. Blood flow is directly proportional to blood pressure and
regulate blood flow into capillary beds.
inversely proportional to resistance.
2. Atherosclerosis is a degenerative vascular disease that
decreases the elasticity of arteries.
Complete an interactive tutorial: > Study Area >
19.3 Capillaries are exchange vessels (pp. 742–744) Interactive Physiology > Cardiovascular System: Factors Affecting
1. Capillaries are microscopic vessels with very thin walls. Most Blood Pressure.
exhibit intercellular clefts, which aid in the exchange between
blood and interstitial fluid. 19.7 Blood pressure decreases as blood flows from arteries
2. The most permeable capillaries are sinusoid capillaries (wide, through capillaries and into veins (pp. 748–750)
19
17 tortuous channels). Fenestrated capillaries with pores are next 1. Systemic blood pressure is highest in the aorta and lowest in the
most permeable. Least permeable are continuous capillaries, venae cavae. The steepest drop in BP occurs in the arterioles,
which lack pores. where resistance is greatest.
3. Most capillary beds consist of a terminal arteriole leading into 2. Arterial BP depends on compliance of the elastic arteries and
capillaries drained by a postcapillary venule. The diameter on how much blood is forced into them. Arterial blood pressure
of the terminal arteriole and upstream arterioles determines is pulsatile, and peaks during systole; this is measured as
the amount of blood flowing through the capillaries. In select systolic pressure. During diastole, as blood is forced distally
capillary beds (e.g., mesenteric capillary beds), vascular shunts in the circulation by the rebound of elastic arteries, arterial BP
(metarteriole–thoroughfare channels) connect the terminal drops to its lowest value, called the diastolic pressure.
arteriole and postcapillary venule at opposite ends of a capillary 3. Pulse pressure is systolic pressure minus diastolic pressure.
bed. In this case, the amount of blood flowing into the true The mean arterial pressure (MAP) 5 diastolic pressure plus
capillaries is regulated by precapillary sphincters. one-third of pulse pressure and is the pressure that keeps blood
19.4 Veins are blood reservoirs that return blood toward moving throughout the cardiac cycle.
the heart (pp. 744–746) 4. Pulse and blood pressure measurements are used to assess
cardiovascular efficiency.
1. Veins have comparatively larger lumens than arteries, and a
5. The pulse is the alternating expansion and recoil of arterial
system of valves prevents backflow of blood.
walls with each heartbeat. Pulse points are also pressure points.
2. Normally most veins are not filled to capacity; for this reason,
6. Blood pressure is routinely measured by the auscultatory method.
they can serve as blood reservoirs.
Normal BP in adults is 120/80 mm Hg (systolic/diastolic).
7. Low capillary pressure (35 to 17 mm Hg) protects the delicate
capillaries from rupture while still allowing adequate exchange
across the capillary walls.
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8. Venous pressure is nonpulsatile and low (declining to zero) maintain MAP and redistribute blood during exercise and
because of the cumulative effects of resistance. Venous valves, thermoregulation.
large lumens, functional adaptations (muscular and respiratory 2. Autoregulation involves myogenic controls that maintain flow
pumps), and sympathetic nervous system activity promote despite changes in blood pressure, and local chemical factors.
venous return. Vasodilators include increased CO2, H+, and nitric oxide.
Decreased O2 concentrations also cause vasodilation. Other
19.8 Blood pressure is regulated by short- and factors, including endothelins, decrease blood flow.
long-term controls (pp. 750–757) 3. In most instances, autoregulation is controlled by the
1. Blood pressure varies directly with cardiac output, total accumulation of local metabolites and the lack of oxygen.
peripheral resistance (TPR), and blood volume. Vessel diameter However, autoregulation in the brain is controlled primarily by
is the major factor determining resistance, and small changes in a drop in pH and by myogenic mechanisms; and pulmonary
the diameter of vessels (chiefly arterioles) significantly affect circuit vessels dilate in response to high levels of oxygen.
blood pressure.
2. BP is regulated by autonomic neural reflexes involving 19.10 Slow blood flow through capillaries promotes
baroreceptors or chemoreceptors, the cardiovascular center diffusion of nutrients and gases, and bulk flow of
(a medullary center that includes the cardiac and vasomotor fluids (pp. 762–766)
centers), and autonomic fibers to the heart and vascular smooth 1. Blood flows fastest where the cross-sectional area of the
muscle. vascular bed is least (aorta), and slowest where the total
3. Activation of the receptors by falling BP (and to a lesser cross-sectional area is greatest (capillaries). The slow flow in
extent by a rise in blood CO2, or falling blood pH or O2 levels) capillaries allows time for nutrient-waste exchanges.
stimulates the vasomotor center to increase vasoconstriction 2. Nutrients, gases, and other solutes smaller than plasma proteins
and the cardioacceleratory center to increase heart rate and cross the capillary wall by diffusion; larger molecules are
contractility. Rising BP inhibits the vasomotor center (permitting actively transported via pinocytotic vesicles. Water-soluble
vasodilation) and activates the cardioinhibitory center. substances move through the clefts or fenestrations; fat-soluble
4. Higher brain centers (cerebrum and hypothalamus) may modify substances pass through the lipid portion of the endothelial cell
neural controls of BP via medullary centers. membrane.
5. Hormones that increase BP by promoting vasoconstriction 3. Bulk flow of fluids at capillary beds determines the distribution
include epinephrine and NE (these also increase heart rate and of fluids between the bloodstream and the interstitial space. It
contractility), ADH, and angiotensin II (generated in response reflects the relative effects of hydrostatic and osmotic pressures
to renin release by kidney cells). Atrial natriuretic peptide is a acting at the capillary (outward minus inward pressures). In
hormone that promotes vasodilation and also causes a decline general, fluid flows out of the capillary bed at the arteriolar end
in blood volume. and reenters the capillary blood at the venule end.
6. The kidneys regulate blood pressure by regulating blood 4. Lymphatic vessels collect the small net loss of fluid and protein
volume. Rising BP directly enhances filtrate formation and into the interstitial space and return it to the cardiovascular
fluid losses in urine; falling BP causes the kidneys to retain system.
more water, increasing blood volume. 5. Edema is an abnormal accumulation of fluid in the interstitial
7. Indirect renal regulation of blood volume involves the renin- space as a result of imbalances in pressures that drive bulk flow
angiotensin-aldosterone mechanism, a hormonal mechanism. or a block of lymphatic drainage.
When BP falls, the kidneys release renin, which triggers the
formation of angiotensin II. Angiotensin II causes (1) release
PART 3
of aldosterone, stimulating salt and water retention,
(2) vasoconstriction, (3) release of ADH, and (4) thirst. CIRCULATORY PATHWAYS: 19
17
8. Chronic hypertension (high blood pressure) is persistent BP BLOOD VESSELS OF THE BODY
readings of 140/90 or higher. It indicates increased total peripheral 1. The pulmonary circulation transports O2-poor, CO2-laden blood
resistance, which strains the heart and promotes vascular to the lungs for oxygenation and carbon dioxide unloading.
complications of other organs, particularly the eyes and kidneys. Blood returning to the right atrium of the heart is pumped by
It is a major cause of myocardial infarction, stroke, and renal the right ventricle to the lungs via the pulmonary trunk. Blood
disease. Risk factors are high-fat, high-salt diet, obesity, diabetes issuing from the lungs is returned to the left atrium by the
mellitus, advanced age, smoking, stress, and being a member of pulmonary veins. (See Table 19.3 and Figure 19.21.)
the black race or a family with a history of hypertension. 2. The systemic circulation transports oxygenated blood from the
9. Hypotension, or low blood pressure (below 90/60 mm Hg), is left ventricle to all body tissues via the aorta and its branches.
rarely a problem except in circulatory shock. Venous blood returning from the systemic circuit is delivered to
10. Circulatory shock occurs when blood perfusion of body tissues the right atrium via the venae cavae.
is inadequate. Most cases of shock reflect low blood volume 3. All arteries are deep, while veins are both deep and superficial.
(hypovolemic shock), abnormal vasodilation (vascular shock), Superficial veins tend to have numerous interconnections. Dural
or pump failure (cardiogenic shock). venous sinuses and the hepatic portal circulation are unique
venous drainage patterns.
19.9 Intrinsic and extrinsic controls determine blood flow
through tissues (pp. 757–762) 19.8 The vessels of the systemic circulation transport
1. Intrinsic controls (autoregulation) involve local adjustment blood to all body tissues (pp. 767–791)
of blood flow to individual organs based on their immediate 1. Tables 19.3 to 19.13 and Figures 19.22 to 19.32 illustrate and
requirements. Extrinsic controls (nerves and hormones) describe vessels of the systemic circulation.
M19_MARI0853_11_GE_C19.indd 795 03/07/18 2:06 PM

Developmental Aspects of Blood Vessels (p. 791) 3. Blood pressure is low in infants and rises to adult values.
1. The fetal vasculature develops from embryonic blood islands and Age-related vascular problems include varicose veins,
mesenchyme and functions in blood delivery by the fourth week. hypertension, and atherosclerosis. Hypertension and associated
2. Fetal circulation differs from circulation after birth. The atherosclerosis are the most important causes of cardiovascular
pulmonary and hepatic shunts and special umbilical vessels are disease in the aged.
normally occluded shortly after birth.
REVIEW QUESTIONS To access additional practice questions using your smartphone, tablet,
or computer: > Study Area > Practice Tests & Quizzes
Level 1 Remember/Understand 11. The only blood vessels branching off the ascending aorta
(Some questions have more than one correct answer. Select the best are the (a) common carotid arteries, (b) coronary arteries,
answer or answers from the choices given.) (c) subclavian arteries, (d) common iliac arteries.
1. Which of the following is true about veins? (a) Venous valves are Level 2 Apply/Analyze
formed from the tunica media. (b) Up to 35% of total body blood 12. Tracing the blood from the heart to the left foot, we find that
is in venous circulation at any given time. (c) Veins have a small blood passes through the aortic arch, the thoracic aorta, the
lumen in relation to the thickness of the vessel wall. (d) Veins are abdominal aorta, the left common iliac artery, the external
called capacitance vessels or blood reservoirs. iliac artery, the popliteal artery, and the posterior tibial artery
2. Total peripheral resistance (a) is inversely proportional to the to arrive at the left foot. Which artery is missing from this
length of the vascular bed, (b) increases in anemia, (c) decreases sequence? (a) internal iliac, (b) axillary, (c) subclavian,
in polycythemia, (d) is inversely related to the diameter of the (d) femoral.
arterioles. 13. How is the anatomy of capillaries and capillary beds well suited
3. Which of the following can lead to increased stroke volume and to their function?
cardiac output? (a) decreased venous return, (b) stimulation of 14. (a) Define blood pressure. Differentiate between systolic and
the cardioinhibitory center, (c) an increase in ANP secretion, diastolic blood pressure. (b) What is the normal blood pressure
(d) increased activity of the respiratory pump. value for an adult?
4. Arteriolar blood pressure increases in response to all but which 15. Describe the short-term hormonal controls regulating blood
of the following? (a) increasing stroke volume, (b) increasing pressure.
heart rate, (c) rising blood volume, (d) falling blood volume. 16. How are nutrients, wastes, and respiratory gases transported to
5. Which of the following would not result in the dilation of the and from the blood and tissue spaces?
terminal arterioles and upstream arterioles in systemic capillary 17. (a) What blood vessels contribute to the formation of the
beds? (a) a decrease in local tissue O2 content, (b) an increase hepatic portal circulation? (b) Why is a portal circulation a
in local tissue CO2, (c) a local increase in histamine, (d) a local “strange” circulation?
increase in pH. 18. Physiologists often consider capillaries and postcapillary
6. Sinusoid capillaries (a) have large fenestrations and venules together. (a) What functions do these vessels share?
intercellular clefts, (b) occur in liver and spleen, (c) have larger (b) Structurally, how do they differ?
lumens than other capillaries, (d) all of these.
7. The baroreceptors in the carotid sinus and aortic arch are Level 3 Evaluate/Synthesize
19
17 sensitive to (a) a decrease in CO2, (b) changes in arterial 19. Distinguish between elastic arteries, muscular arteries, and
pressure, (c) a decrease in O2, (d) all of these. arterioles relative to location, histology, and functional
8. Blood draining from the brain enters the (a) coronary sinus, adaptations.
(b) cephalic vein, (c) dural venous sinus, (d) inferior vena cava. 20. Write an equation showing the relationship between total
9. Blood flow in the capillaries is steady despite the rhythmic peripheral resistance, blood flow, and blood pressure.
pumping of the heart because of the (a) elasticity of the large 21. Explain the reasons for the observed changes in blood flow
arteries, (b) small diameter of capillaries, (c) thin walls of the velocity in the different regions of the circulation.
veins, (d) venous valves. 22. Excessive sweating during strenuous exercise can cause
10. Using the letters from column B, match the artery descriptions in a decrease in blood volume and pressure. Which neural
column A. (Note that some require more than a single choice.) mechanisms will be activated to restore blood volume and
Column A Column B pressure?
____ (1) unpaired branch of (a) right common carotid 23. Describe neural and chemical (both systemic and local) effects
abdominal aorta (b) superior mesenteric exerted on the blood vessels when you are fleeing from a mugger.
____ (2) second branch of (c) left common carotid (Be careful, this is more involved than it appears at first glance.)
aortic arch (d) external iliac 24. A 60-year-old man is unable to walk more than 100 yards without
____ (3) branch of internal (e) inferior mesenteric experiencing severe pain in his left leg; the pain is relieved by
carotid (f) superficial temporal resting for 5–10 minutes. He is told that the arteries of his leg are
____ (4) branch of external (g) celiac trunk becoming occluded with fatty material and is advised to have the
carotid (h) facial sympathetic nerves serving that body region severed. Explain
____ (5) origin of femoral (i) ophthalmic how such surgery might help to relieve this man’s problem.
arteries (j) internal iliac 25. Your friend Jillian, who knows little about science, is reading a
magazine article about a patient who had an “aneurysm at the base
M19_MARI0853_11_GE_C19.indd 796 03/07/18 2:06 PM

of his brain that suddenly grew much larger.” The surgeons’ first
goal was to “keep it from rupturing,” and the second goal was to CLINICAL CASE STUDY
“relieve the pressure on the brain stem and cranial nerves.” The
surgeons were able to “replace the aneurysm with a section of 35-Year-Old Male with Hypovolemic Shock
plastic tubing,” so the patient recovered. Jillian asks you what all Mr. Clark, a 35-year-
this means. Explain. (Hint: Check this chapter’s Related Clinical old male, met with
Terms, p. 791.) an accident as he lost
26. The Agawam High School band is playing some lively marches control of his motor-
while the coaches are giving pep talks to their respective cycle on a wet hilly
football squads. Although it is September, it is unseasonably road. Upon arrival at
hot (88°F/31°C) and the band uniforms are wool. Suddenly the scene, paramedics
Ryan, the tuba player, becomes light-headed and faints. Explain quickly stabilize Mr.
his fainting in terms of vascular events. Clark, and prepare to
27. When we are cold or the external temperature is low, most take him to the nearest
venous blood returning from the distal part of the arm travels hospital. On their way,
in the deep veins where it picks up heat (by countercurrent they phone to inform
exchange) from the nearby brachial artery en route. However, the attending nurse
when we are hot, and especially during exercise, venous return that the patient’s distal
from the distal arm travels in the superficial veins and those left femur is fractured,
veins tend to bulge superficially in a person who is working and his popliteal artery
out. Explain why venous return takes a different route in the is severed, but his vital
second situation. signs are good, and
28. Edema is a common clinical problem. On your first day of a his blood pressure is
clinical rotation, you encounter four patients who have edema for 110/90 mm Hg.
different reasons. Your challenge is to explain the edema in terms Mr. Clark’s skin is found to be cyanotic and cool upon
of either an increase or a decrease in one of the four pressures his arrival at the hospital. His blood pressure has dropped to
that causes bulk flow (see Focus Figure 19.1 on pp. 764–765). 85/55 mm Hg, his heart rate is 115 beats/min, and his pulse
(1) First you encounter Mrs. Taylor in the medical unit awaiting is weak. The physician diagnoses Mr. Clark with second
a liver transplant. What is the connection between liver failure stage of hypovolemic shock, and estimates that he has lost
and her edema? about 20% of his blood volume.
(2) Next in the obstetric ward, Mrs. So is experiencing premature 1. Where in the body is the popliteal artery located, and
labor and has edema in her legs. Which bulk flow pressures which body parts does it supply with blood?
might be altered here?
(3) In emergency, Mr. Herrera is in anaphylactic shock. His capil- 2. If Mr. Clark has lost 20% of his blood volume, how much
laries have become leaky, allowing plasma proteins that are blood (in liters) has he actually lost?
normally kept inside the blood vessels to escape into the inter- 3. ✚ NCLEX-STYLE When a patient experiences a significant
stitial fluid. Which of the bulk flow pressures is altered in this loss of blood or body fluids, it results in a drop in blood
case and in what direction is the change? volume and also blood pressure. The decrease in Mr.
(4) Finally, in oncology Mrs. O’Leary is recovering from breast Clark’s blood pressure will be detected by the baroreceptors
cancer surgery. Her right breast and all of her axillary lymph in the carotid sinuses and aortic arch. These baroreceptors
nodes were removed. Unfortunately, this severed most of will send impulses to the medulla oblongata, which will: 19
17
the lymphatic vessels draining her right arm. You notice that a. Activate the cardioinhibitory center
this arm is quite edematous. Why? Mrs. O’Leary is given a b. Activate the cardioacceleratory center
compression sleeve to wear on this arm to help relieve the c. Inhibit the vasomotor center
edema. Which of the bulk flow pressures will be altered by the d. Inhibit the respiratory centers
compression sleeve? 4. Mr. Clark’s cyanotic, cool skin is a result of stimulation of
his vasomotor center. Why will stimulation of this brain
center cause these symptoms, and how will it assist to
compensate for the decreased blood pressure?
5. ✚ NCLEX-STYLE Although Mr. Clark’s pulse is weak, he
has an increased heart rate. His increased heart rate is
brought about by:
a. Sympathetic innervation of the AV node
b. Parasympathetic innervation of the SA node
c. Sympathetic innervation of the myocardium
d. Sympathetic innervation of the SA node
6. In hypovolemic shock, antidiuretic hormone (ADH) plays
a role in both the short-term and long-term responses to
the loss of blood or body fluids. Explain the difference
between these two responses induced by ADH.
M19_MARI0853_11_GE_C19.indd 797 03/07/18 2:06 PM

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19 The Cardiovascular System:

In this chapter, you will learn that

Part 1 Blood Vessel Part 2 Physiology Part 3 Circulatory

by examining by asking by exploring

19.1 Structure of 19.11 Principal vessels of

19.7 What is blood and finally, exploring

19.5 Anastomoses 19.8 How is blood

Watch a video to learn

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Check Your Understanding

M19_MARI0853_11_GE_C19.indd 741 03/07/18 2:05 PM

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(a) Continuous capillary

• Abundant in skin, muscles, lungs, and CNS. Red blood

(b) Fenestrated capillary

Fenestrated capillaries have large fenestrations (pores) Pinocytotic

(c) Sinusoid capillary

• Occur in liver, bone marrow, spleen, and adrenal medulla.

Figure 19.3 Capillary structure.

M19_MARI0853_11_GE_C19.indd 743 03/07/18 2:05 PM

Capillary Beds Vascular shunt

Veins are blood reservoirs that

M19_MARI0853_11_GE_C19.indd 744 03/07/18 2:05 PM

Venules is low. However, the low-pressure condition demands several

M19_MARI0853_11_GE_C19.indd 745 03/07/18 2:05 PM

M19_MARI0853_11_GE_C19.indd 746 03/07/18 2:05 PM

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aorta as aortic pressure reaches its peak. Indeed, if the aorta

0 Clinical Monitoring of Circulatory Efficiency

measuring pulse and blood pressure. These values, along with

measurements of respiratory rate and body temperature, are

referred to collectively as the body’s vital signs. Let’s examine

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Check Your Understanding

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Stroke volume Heart rate Diameter of Blood viscosity Blood vessel

Cardiac output Total peripheral resistance

Mean arterial pressure (MAP)

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3 Impulses from baroreceptors activity, reducing heart rate

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Table 19.2 Effects of Selected Hormones on Blood Pressure

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Direct renal mechanism Indirect renal mechanism (renin-angiotensin-aldosterone)

Filtration by kidneys Angiotensinogen

ADH release by Thirst via Vasoconstriction; total

M19_MARI0853_11_GE_C19.indd 754 03/07/18 2:06 PM

Conservation Blood volume Blood pH

Blood Baroreceptors Chemoreceptors

Venous Activation of vasomotor and cardio-

Diameter of Blood Blood vessel

Cardiac output Total peripheral resistance

Figure 19.13 Factors that increase MAP.

M19_MARI0853_11_GE_C19.indd 755 03/07/18 2:06 PM

and interfere with the ability of endothelial cells to induce

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You can compare blood flow autoregulation to water use in

Table 19.3 (continued)

Table 19.4 (continued)

Table 19.5 (continued)

Table 19.6 (continued)