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Look for these other volumes in the Mosby Physiology Series:

Blaustein, Kao, & Matteson: CELLULAR PHYSIOLOGY AND NEUROPHYSIOLOGY

Cloutier: RESPIRATORY PHYSIOLOGY

Hudnall: HEMATOLOGY: A PATHOPHYSIOLOGIC APPROACH

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Koeppen & Stanton: RENAL PHYSIOLOGY

White, Harrison, & Mehlmann: ENDOCRINE AND REPRODUCTIVE PHYSIOLOGY

CARDIOVASCULAR
PHYSIOLOGY
11TH EDITION

Achilles J. Pappano, PhD

Professor Emeritus
Department of Cell Biology and Calhoun Cardiology Center
University of Connecticut Health Center
Farmington, Connecticut
Withrow Gil Wier, PhD
Professor Emeritus
Department of Physiology
University of Maryland School of Medicine
Baltimore, MD
1600 John F. Kennedy Blvd.
Ste 1800
Philadelphia, PA 19103-2899

CARDIOVASCULAR PHYSIOLOGY, ELEVENTH EDITION ISBN: 978-0-323-59484-4

Copyright © 2019 Elsevier Inc. All Rights Reserved.

Previous editions copyrighted 2013, 2007, 2001, 1997

No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechan-
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This book and the individual contributions contained in it are protected under copyright by the Publisher (other
than as may be noted herein).

Notices

Practitioners and researchers must always rely on their own experience and knowledge in evaluating and
using any information, methods, compounds or experiments described herein. Because of rapid advances in
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Printed in the United States of America

Last digit is the print number: 9 8 7 6 5 4 3 2 1

 D E D I C AT I O N

To Robert M. Berne and Matthew N. Levy,

whose research and scholarship in cardiovascular physiology have enriched
and inspired generations of students and colleagues
P R E FA C E
We believe that physiology is the backbone of clinical med- coronary blood flows have received particular emphasis.
icine. In the clinic, the emergency room, the intensive care The theory of cardiac excitation–contraction coupling has
unit, or the surgical suite, physiological principles are the been extensively updated, particularly with respect to new
basis for action. But we also find great intellectual satisfac- understanding of the roles of intracellular calcium ions.
tion in the science of physiology as the means to explain the Whenever available, physiological data from humans have
elegant mechanisms of our bodies. In the eleventh edition been included. Some old figures have been deleted and
of Berne and Levy’s classic monograph on cardiovascular many new figures have been added to aid comprehension
physiology, we have tried to convey both ideas. of the text. Selected references appear at the end of each
Physiology serves as a foundation that students of chapter. The scientific articles included were chosen for
medicine must comprehend before they can understand their depth, clarity, and appropriateness.
the derangements caused by pathology. This text of car- Throughout the book, italics are used to emphasize
diovascular physiology emphasizes general concepts and important facts and concepts, and boldface type is used
regulatory mechanisms. To present the various regulatory for new terms and definitions. Each chapter begins with a
mechanisms clearly, the component parts of the system are list of objectives and ends with a summary to highlight key
first discussed individually. Then, the last chapter describes points. Case histories with multiple-choice questions are
how various individual components of the cardiovascu- provided to help in review and to indicate clinical relevance
lar system are coordinated. The examples describe how of the material. The correct answers and brief explanations
the body responds to two important stresses—exercise for them appear in Appendix A. A comprehensive review
and hemorrhage. Selected pathophysiological examples examination, with explanations of the correct answers, has
of abnormal function are included to illustrate and clarify been added as Appendix B.
normal physiological processes. These examples are dis- We thank our readers for their constructive comments.
tributed throughout the text and are identified by colored Thanks are also due to the numerous investigators and
boxes with the heading “Clinical Box.” publishers who have granted permission to use illustra-
The text incorporates the learning objectives for cardio- tions from their publications. In most cases these illustra-
vascular physiology of the American Physiological Society, tions have been altered somewhat to increase their didactic
except for hemostasis and coagulation. These last-named utility. In some cases, unpublished data from investiga-
topics are found in hematology books. The book has been tions by Robert Berne and Matthew Levy and the current
updated and revised extensively. The relation between authors have been presented.
pressure–volume loops and cardiac function curves, newer
aspects of endothelium function, myocardial metabo- Achilles J. Pappano
lism and its relation to oxygen consumption and cardiac W. Gil Wier
energetics, and the regulation of peripheral, cerebral, and

vi
 Overview of the Circulation and Blood

OBJECTIVES
1. Describe the general structure of the cardiovascular 4. Indicate the pressure changes and pathways of blood
system. flow throughout the vasculature.
2. Compare the compositions and functions of the blood 5. Describe the constituents of the blood and explain the
vessels. functions of the cellula elements of blood.
3. Compare the relationship of the vascular cross-sec- 6. Know the importance 0f blooa. group matching before
tional area to the velocity of blood flow in the various blood transfusions.
vascular segments.

The circulatory, endocrine, and nervous systems constitute consists of two pumps in series: the right ventricle to propel
the principal coordinating and integrating systems of the blood througH the lungs for exchange of 0 2 and CO 2 (the
body. Whereas the nervous system is primarily concerned pulmonary circulation) and the left ventricle to propel
with communication and the endocrine glands with reg- blood to all other tissues of the body (the systemic cir-
ulation of certain body functions, the circulatory system culation). The total flow of blood out of the left ventricle
serves to transport and distribute essential substancF to is known as the cardiac output (CO). The rhythmic con-
the tissues and to remove metabolic byproducts. The circu- traction of the heart is an intrinsic property of the heart
latory system also shares in such homeostatic mechanisms whose sinoatrial node pacemaker generates action paten -
as regulation of body temperature, humoral communi- tials spontaneously (see Chapter 3). These action potentials
cation throughout the body, and adjustments of 0 2 and are propagated in an orderly manner through the organ to
nutrient supply in different physiologica1 states. trigger contraction and to produce the currents detected in
the electrocardiogram (see Chapter 3).
Unidirectional flow through the heart is achieved
by the appropriate arrangement of effective flap valves.
The cardiovascular system accomp ishes these functions Although the cardiac output is intermittent, continuous
with a pump (see Chapter 4), a sec·es of distributing and flow to the periphery occurs by distention of the aorta and
collecting tubes (see Chapter 7), and an extensive system its branches during ventricular contraction (systole) and
of thin vessels that permit rapid exchange between the tis- elastic recoil of the walls of the large arteries that propel the
sues and the vascular channels (see Chapter 8). The pri- blood forward during ventricular relaxation (diastole).
mary purpose of this text is to discuss the function of the Blood moves rapidly through the aorta and its arterial
components of the vascular system and the control mecha- branches (see Chapter 7). The branches become narrower
nisms (with their checks and balances) that are responsible and their walls become thinner and change histologi-
for alteration of blood distribution necessary to meet the cally toward the periphery. From the aorta, a predomi-
changing requirements of different tissues in response to a nantly elastic structure, the peripheral arteries become
wide spectrum of physiological (see Chapters 9 and 10) and more muscular until the muscular layer predominates
pathological (see Chapter 13 ) conditions. at the arterioles (Fig. 1.2 ).
Before one considers the function of the parts of the In the large arteries, frictional resistance is relatively
circulatory system in detail, it is useful to consider it as a small, and mean pressure throughout the system of large
whole in a purely descriptive sense (Fig. 1. 1). The heart arteries is only slightly less than in the aorta. The small

1
2 CHAPTER 1 Overview of the Circulation and Blood

Veins Arteries In addition to a sharp reduction in pressure across the

arterioles, there is also a change from pulsatile to steady
Venules flow as pressure continues to decline from the arterial to
Head and neck
Capillaries Arterioles the venous end of the capillaries (see Fig. 1.3). The pulsatile
arteries
arterial blood flow, caused by the phasic cardiac ejection, is
Pulmonary veins damped at the capillaries by the combination of distensibility
Arm arteries
of the large arteries and frictional resistance in the arterioles.
Bronchial arteries
Pulmonary CLINICAL BOX
artery
In a patient with hyperthyroidism (Graves disease), the
basal metabolism is elevated and is often associated
Right atrium Left atrium Aorta with arteriolar vasodilation. This reduction in arteriolar
Left ventricle resistance diminishes the dampening effect on the pul-
Coronary satile arterial pressure and is manifested as pulsatile
Venae cavae
arteries flow in the capillaries, as observed in the fingernail beds
Right Splenic of patients with this ailment.
ventricle artery
Trunk arteries
Many capillaries arise from each arteriole to form
Hepatic the microcirculation (see Chapter 8), so that the total
vein Hepatic artery
cross-sectional area of the capillary bed is very large, despite
the fact that the cross-sectional area of each capillary is less
Portal vein
than that of each arteriole. As a result, blood flow velocity
Peritubular Renal
becomes quite slow in the capillaries (see Fig. 1.3), anal-
capillaries Mesenteric arteries ogous to the decrease in velocity of flow seen at the wide
arteries
regions of a river. Conditions in the capillaries are ideal for
Afferent the exchange of diffusible substances between blood and
Efferent arterioles
Glomeruli arterioles tissue, because the capillaries are short tubes whose walls
are only one cell thick and because flow velocity is low.
On its return to the heart from the capillaries, blood
Pelvic arteries passes through venules and then through veins of increas-
Leg arteries
ing size with a progressive decrease in pressure until the
blood reaches the vena cava (see Fig. 1.3). As the heart is
Fig. 1.1 Schematic diagram of the parallel and series arrange- approached, the number of veins decreases, the thickness
ment of the vessels composing the circulatory system. The cap- and composition of the vein walls change (see Fig. 1.2), the
illary beds are represented by thin lines connecting the arteries
total cross-sectional area of the venous channels dimin-
(on the right) with the veins (on the left). The crescent-shaped
thickenings proximal to the capillary beds represent the arteri-
ishes, and the velocity of blood flow increases (see Fig. 1.3).
oles (resistance vessels). (Redrawn from Green, H. D. (1944). In Note that the velocity of blood flow and the cross-sectional
O. Glasser (Ed.). Medical physics (Vol 1); Chicago: Mosby-Year area at each level of the vasculature are essentially mirror
Book.) images of each other (see Fig. 1.3).
Data indicate that between the aorta and the capillar-
ies the total cross-sectional area increases about 500-fold
arteries and arterioles serve to regulate flow to individ- (see Fig. 1.3). The volume of blood in the systemic vas-
ual tissues by varying their resistance to flow. The small cular system (Table 1.1) is greatest in the veins and small
arteries offer moderate resistance to blood flow, and this veins (64%). Of the total blood volume only about 6% is
resistance reaches a maximal level in the arterioles, some- in the capillaries and 14% in the aorta, arteries, and arteri-
times referred to as the stopcocks of the vascular system. oles. In contrast, blood volume in the pulmonary vascular
Hence the pressure drop is significant and is greatest in the bed is about equal between arteries and capillaries; venous
small arteries and in the arterioles (Fig. 1.3). Adjustments vessels display a slightly larger percentage of pulmonary
in the degree of contraction of the circular muscle of blood volume. The cross-sectional area of the venae cavae
these small vessels permit regulation of tissue blood flow is larger than that of the aorta. Therefore the velocity of
and aid in the control of arterial blood pressure (see flow is slower in the venae cavae than that in the aorta
Chapter 9). (see Fig. 1.3).
 CHAPTER 1 Overview of the Circulation and Blood 3

Macrovessels 10 mm Microvessels 20 µm
Terminal
Aorta Artery Vein Vena cava Arteriole arteriole Capillary Venule
Diameter 25 mm 4 mm 5 mm 30 mm 30 µm 10 µm 8 µm 20 µm

1.5
Wall thickness 2 mm 1 mm 0.5 mm mm 6 µm 2 µm 0.5 µm 1 µm

Endothelium

Elastic tissue

Smooth muscle

Fibrous tissue

Fig. 1.2 Internal diameter, wall thickness, and relative amounts of the principal components of the vessel
walls of the various blood vessels that compose the circulatory system. Cross sections of the vessels are not
drawn to scale because of the huge range from aorta and venae cavae to capillary. (Redrawn from Burton, A.
C. (1954). Relation of structure to function of the tissues of the wall of blood vessels. Physiological Reviews,
34(4), 619–642.)

Blood entering the right ventricle via the right atrium is Furthermore, blood transports other substances, such as
pumped through the pulmonary arterial system at a mean hormones, white blood cells, and platelets, from their sites
pressure about one-seventh that in the systemic arteries. of production to their sites of action. Blood also aids in
The blood then passes through the lung capillaries, where the distribution of fluids, solutes, and heat. Hence blood
CO2 is released and O2 taken up. The O2-rich blood returns contributes to homeostasis, the maintenance of a constant
via the four pulmonary veins to the left atrium and ventri- internal environment.
cle to complete the cycle. Thus in the normal intact circula- A fundamental characteristic of normal operation of
tion the total volume of blood is constant, and an increase the cardiovascular system is the maintenance of a relatively
in the volume of blood in one area must be accompanied constant mean (average) blood pressure within the large
by a decrease in another. However, the distribution of the arteries. The difference between mean arterial pressure (Pa )
circulating blood to the different body organs is deter- and the pressure in the right atrium (Pra) provides the
mined by the output of the left ventricle and by the con- driving force for flow through the resistance (R) of blood
tractile state of the arterioles (resistance vessels) of these vessels of the individual tissues. Thus when the circulatory
organs (see Chapters 9 and 10). In turn, the cardiac output system is in steady-state, total flow of blood from the heart
is controlled by the rate of heartbeat, cardiac contractility, (cardiac output, CO) equals total flow of blood returning
venous return, and arterial resistance. The circulatory sys- to the heart. The relation among these variables is described
tem is composed of conduits arranged in series and in par- in the following hydraulic equation:
allel (see Fig. 1.1).
It is evident that the systemic and pulmonary vascular Pa − Pra = CO × R (1.1)
systems are composed of many blood vessels arranged in
series and parallel, with respect to blood flow. The total The cardiovascular system, together with neural, renal,
resistance to blood flow of the systemic blood vessels is and endocrine systems, maintains Pa at a relatively con-
known as the total peripheral resistance (TPR), and the total stant level, despite the large variations in cardiac output
resistance of the pulmonary vessels is known as the total pul- and peripheral resistance that are required in daily life. If
monary resistance. Total peripheral resistance and cardiac the Pa is maintained at its normal level under all circum-
output determine the mean pressure in the large arteries, stances, then each individual tissue will be able to obtain the
through the hydraulic resistance equation (see Chapter 7). necessary blood flow required to sustain its functions. Because
The main function of the circulating blood is to carry blood flow to the brain and the heart cannot be interrupted
O2 and nutrients to the various tissues in the body and for even a few seconds without endangering life, maintenance
to remove CO2 and waste products from those tissues. of the Pa is a critical function of the cardiovascular system.
4 CHAPTER 1 Overview of the Circulation and Blood

120

Pressure (mmHg)
80

40 (Pulmonary
artery)

Total cross-section area (cm2) Blood velocity (cm/s)

40

Aorta
23 Vena cava
20 (mean) 15

1000

100 Vena cava

7
10 Aorta
Fig. 1.3 Phasic pressure, velocity of flow, and cross-sectional 4
area of the systemic circulation. The important features are the 0
major pressure drop across the small arteries and arterioles,
the inverse relationship between blood flow velocity and cross-

va
sectional area, and the maximal cross-sectional area and min-

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BLOOD
from these stem cells. Most of these immature cells develop
Blood consists of red blood cells, white blood cells, and into various forms of mature cells, such as erythrocytes,
platelets suspended in a complex solution (plasma) of var- monocytes, megakaryocytes, and lymphocytes. The
ious salts, proteins, carbohydrates, lipids, and gases. The erythrocytes lose their nuclei before they enter the circula-
circulating blood volume accounts for about 7% of the tion, and their average life span is 120 days. Approximately
body weight. Approximately 55% of the blood is plasma; 5 million erythrocytes are present per microliter of blood.
the protein content is 7 g/dL (about 4 g/dL of albumin and However, a small fraction of the pluripotential stem cells
3 g/dL of plasma globulins). remains in the undifferentiated state.
Hemoglobin (about 15 g/dL of blood) is the main
Erythrocytes protein in the erythrocytes. Hemoglobin consists of
The erythrocytes (red blood cells) are flexible, biconcave heme, an iron-containing tetrapyrrole. Heme is linked
disks that transport oxygen to the body tissues (Fig. 1.4). to globin, a protein composed of four polypeptide chains
Mammalian erythrocytes are unusual in that they lack a (two α and two β chains in the normal adult). The iron
nucleus. The average erythrocyte is 7 μm in diameter, and moiety of hemoglobin binds loosely and reversibly to O2
these cells arise from pluripotential stem cells in the bone to form oxyhemoglobin. The affinity of hemoglobin for
marrow. All of the cells in the circulating blood are derived O2 is a steep function of the partial pressure of O2 (Po2)
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