Cardiovascular physiology

305 Pages·2013·13.169 MB·English

by  Pappano A.J., Wier W.G.

Cardiovascular
Physiology
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Cardiovascular
Physiology
T E N T H E D I T I O N
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
Department of Physiology
University of Maryland School of Medicine
Baltimore, Maryland
1600 John F. Kennedy Blvd.
Ste 1800
Philadelphia, PA 19103-2899
CARDIOVASCULAR PHYSIOLOGY ISBN: 978-0-323-08697-4
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Library of Congress Cataloging-in-Publication Data
Pappano, Achilles J.
Cardiovascular physiology / Achilles J. Pappano, Withrow Gil Wier. -- 10th ed.
p. ; cm. -- (Mosby physiology monograph series)
Rev. ed. of: Cardiovascular physiology / Matthew N. Levy, Achilles J. Pappano. 9th ed. c2007.
Includes bibliographical references and index.
ISBN 978-0-323-08697-4 (pbk.)
I. Wier, Withrow Gil. II. Levy, Matthew N., 1922- Cardiovascular physiology. III. Title. IV. Series: Mosby
physiology monograph series.
[DNLM: 1. Cardiovascular Physiological Phenomena. WG 102]
612.1--dc23 2012032909
Senior Content Strategist: Elyse O’Grady
Content Coordinator: Lee Hood
Publishing Services Managers: Rajendrababu Hemamalini and Anne Altepeter
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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
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 F A C E
W
e believe that physiology is the backbone of endothelium function, myocardial metabolism and its
clinical medicine. In the clinic, the emergency room, relation to oxygen consumption and cardiac energet-
the intensive care unit, or the surgical suite, physiolog- ics, and the regulation of peripheral and coronary
ical principles are the basis for action. But we also find blood flows have received particular emphasis. When-
great intellectual satisfaction in the science of physiol- ever available, physiological data from humans have
ogy as the means to explain the elegant mechanisms of been included. Some old figures have been deleted and
our bodies. In the tenth edition of Berne and Levy’s many new figures have been added to aid comprehen-
classic monograph on cardiovascular physiology, we sion of the text. Selected references appear at the end
have tried to convey both ideas. of each chapter. The scientific articles included were
Physiology serves as a foundation that students of chosen for their depth, clarity, and appropriateness.
medicine must comprehend before they can under- Throughout the book, italics are used to emphasize
stand the derangements caused by pathology. This text important facts and concepts, and boldface type is
of cardiovascular physiology emphasizes general con- used for new terms and definitions. Each chapter
cepts and regulatory mechanisms. To present the vari- begins with a list of objectives and ends with a sum-
ous regulatory mechanisms clearly, the component mary to highlight key points. Case histories with
parts of the system are first discussed individually. multiple-choice questions are provided to help in
Then, the last chapter describes how various individ- review and to indicate clinical relevance of the mate-
ual components of the cardiovascular system are coor- rial. The correct answers and brief explanations for
dinated. The examples describe how the body responds them appear in the appendix.
to two important stresses—exercise and hemorrhage. We thank our readers for their constructive com-
Selected pathophysiological examples of abnormal ments. Thanks are also due to the numerous investiga-
function are included to illustrate and clarify normal tors and publishers who have granted permission to
physiological processes. These examples are distrib- use illustrations from their publications. In most cases
uted throughout the text and are identified by colored these illustrations have been altered somewhat to
boxes with the heading “Clinical Box”. increase their didactic utility. In some cases, unpub-
The text incorporates the learning objectives for lished data from investigations by Robert Berne and
cardiovascular physiology of the American Physiologi- Matthew Levy and the current authors have been
cal Society, except for hemostasis and coagulation. presented.
These last-named topics are found in hematology
books. The book has been updated and revised exten- Achilles J. Pappano
sively. The relation between pressure-volume loops W. Gil Wier
and cardiac function curves, newer aspects of
vi
C O N T E N T S
1 Conduction in Cardiac Fibers Depends on
C H A P T E R Local Circuit Currents 25
Conduction of the Fast Response  25
OVERVIEW OF THE CIRCULATION
Conduction of the Slow Response  27
AND BLOOD 1
. . . . . . . . . . . . . . . . . . . . . . . . .
Cardiac Excitability Depends on the
The Circulatory System 1
Activation and Inactivation of Specific
Blood 5
Currents 27
Erythrocytes  5
Fast Response  27
Leukocytes  6
Slow Response  28
Lymphocytes  7
Effects of Cycle Length  28
Blood Is Divided into Groups by Antigens 
Summary 29
Located on Erythrocytes  7
Case 2-1 29
Summary 9
Case 1-1 9
3
C H A P T E R
2
AUTOMATICITY: NATURAL
C H A P T E R
EXCITATION OF THE HEART 31
. . .
EXCITATION: THE CARDIAC
The Heart Generates Its Own Pacemaking
ACTION POTENTIAL 11
. . . . . . . . . . . . . Activity 31
Cardiac Action Potentials Consist of Several
Sinoatrial Node  32
Phases 11
Ionic Basis of Automaticity  34
The Principal Types of Cardiac Action 
Overdrive Suppression  35
Potentials Are the Slow and Fast 
Types  12 Atrial Conduction  36
Ionic Basis of the Resting   Atrioventricular Conduction  37
Potential  13 Ventricular Conduction  39
The Fast Response Depends Mainly on  An Impulse Can Travel Around a Reentry
Voltage-Dependent Sodium  Loop 41
Channels  15
Afterdepolarizations Lead to Triggered
Ionic Basis of the Slow Response  24 Activity 42
vii
viii CONTENTS
Early Afterdepolarizations  43 The Two Major Heart Sounds Are 
Delayed Afterdepolarizations  43 Produced Mainly by Closure of the 
Cardiac Valves  74
Electrocardiography Displays the Spread of
Cardiac Excitation 44 The Pressure-Volume Relationships in the
Intact Heart 75
Scalar Electrocardiography  44
Passive or Diastolic Pressure-Volume 
Dysrhythmias Occur Frequently and
Relationship  75
Constitute Important Clinical
Problems 47 Active or End-Systolic Pressure-Volume 
Relationship  77
Altered Sinoatrial Rhythms  47
Pressure and Volume during the Cardiac 
Atrioventricular Transmission Blocks  48
Cycle: The P-V Loop  77
Premature Depolarizations  48
Preload and Afterload during the Cardiac 
Ectopic Tachycardias  49 Cycle  77
Fibrillation  49
Contractility  78
Summary 51
The Fick Principle Is Used to Determine
Case 3-3 52 Cardiac Output 79
Summary 89
Case 4-1 90
4
C H A P T E R
THE CARDIAC PUMP 55 5
. . . . . . . . . . . . .
C H A P T E R
The Gross and Microscopic Structures of the
Heart Are Uniquely Designed for Optimal REGULATION OF THE
Function 55
HEARTBEAT 91
. . . . . . . . . . . . . . . . . . . . . . . .
The Myocardial Cell  55
Heart Rate is Controlled Mainly by the
Structure of the Heart: Atria, Ventricles,  Autonomic Nerves 91
and Valves  60
Parasympathetic Pathways  92
The Force of Cardiac Contraction Is
Sympathetic Pathways  93
Determined by Excitation-Contraction
Higher Centers Also Influence Cardiac 
Coupling and the Initial Sarcomere Length
of the Myocardial Cells 63 Performance  97
Excitation-Contraction Coupling Is  Heart Rate Can Be Regulated via the 
Mediated by Calcium  63 Baroreceptor Reflex  97
Mechanics of Cardiac Muscle  65 The Bainbridge Reflex and Atrial 
Receptors Regulate Heart Rate  98
The Sequential Contraction and Relaxation of
Respiration Induces a Common Cardiac 
the Atria and Ventricles Constitute the
Cardiac Cycle 69 Dysrhythmia  99
Ventricular Systole  70 Activation of the Chemoreceptor Reflex 
Affects Heart Rate  101
Echocardiography Reveals Movement of 
the Ventricular Walls and of the  Ventricular Receptor Reflexes Play a 
Valves  73 Minor Role in the Regulation of Heart 
Rate  102
CONTENTS ix
Myocardial Performance Is Regulated Arterial Elasticity Compensates for the
by Intrinsic Mechanisms 102 Intermittent Flow Delivered by the
The Frank-Starling Mechanism Is an  Heart 137
Important Regulator of Myocardial  The Arterial Blood Pressure Is Determined by
Contraction Force  103 Physical and Physiological Factors 140
Changes in Heart Rate Affect Contractile  Mean Arterial Pressure  140
Force  107 Cardiac Output  142
Myocardial Performance Is Regulated by Peripheral Resistance  142
Nervous and Humoral Factors 110
Pulse Pressure  144
Nervous Control  110
Stroke Volume  144
Cardiac Performance Is Also Regulated by 
Arterial Compliance  145
Hormonal Substances  113
Total Peripheral Resistance and Arterial 
Summary 116
Diastolic Pressure  146
Case 5-1 117
The Pressure Curves Change in Arteries at
Different Distances from the Heart 147
Blood Pressure Is Measured by a
6
C H A P T E R Sphygmomanometer in Human
Patients 148
HEMODYNAMICS 119
. . . . . . . . . . . . . . . Summary 150
Velocity of the Bloodstream Depends on Case 7-1 150
Blood Flow and Vascular Area 119
Blood Flow Depends on the Pressure
Gradient 120 8
C H A P T E R
Relationship Between Pressure and Flow
Depends on the Characteristics of the
THE MICROCIRCULATION
Conduits 122
AND LYMPHATICS . 153
. . . . . . . . . . . . .
Resistance to Flow 125
Functional Anatomy 153
Resistances in Series and in Parallel  126
Arterioles Are the Stopcocks of the 
Flow May Be Laminar or Turbulent 127
Circulation  153
Shear Stress on the Vessel Wall 128
Capillaries Permit the Exchange of Water, 
Rheologic Properties of Blood 129 Solutes, and Gases  154
Summary 133 The Law of Laplace Explains How 
Case 6-6 134 Capillaries Can Withstand High 
Intravascular Pressures  155
The Endothelium Plays an Active Role in
7 Regulating the Microcirculation 156
C H A P T E R
The Endothelium is at the Center of Flow-
THE ARTERIAL SYSTEM 135 Initiated Mechanotransduction 157
. . . . . . .
The Hydraulic Filter Converts Pulsatile Flow The Endothelium Plays a Passive Role in
to Steady Flow 135 Transcapillary Exchange 158