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Guyton and Hall Textbook of Medical Physiology
Guyton and Hall Textbook of Medical Physiology
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Twelfth Edition
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To
My Family
For their abundant support, for their patience and
understanding, and for their love
To
Arthur C. Guyton
For his imaginative and innovative research
For his dedication to education
For showing us the excitement and joy of physiology
And for serving as an inspirational role model
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Preface
The first edition of the Textbook of Medical Physiology I have attempted to maintain the same unified orga-
was written by Arthur C. Guyton almost 55 years ago. nization of the text that has been useful to students in
Unlike most major medical textbooks, which often have the past and to ensure that the book is comprehensive
20 or more authors, the first eight editions of the Textbook enough that students will continue to use it during their
of Medical Physiology were written entirely by Dr. Guyton, professional careers.
with each new edition arriving on schedule for nearly 40 My hope is that this textbook conveys the majesty of
years. The Textbook of Medical Physiology, first published the human body and its many functions and that it stim-
in 1956, quickly became the best-selling medical physi- ulates students to study physiology throughout their
ology textbook in the world. Dr. Guyton had a gift for careers. Physiology is the link between the basic sciences
communicating complex ideas in a clear and interesting and medicine. The great beauty of physiology is that it
manner that made studying physiology fun. He wrote the integrates the individual functions of all the body’s differ-
book to help students learn physiology, not to impress his ent cells, tissues, and organs into a functional whole, the
professional colleagues. human body. Indeed, the human body is much more than
I worked closely with Dr. Guyton for almost 30 years the sum of its parts, and life relies upon this total function,
and had the privilege of writing parts of the 9th and 10th not just on the function of individual body parts in isola-
editions. After Dr. Guyton’s tragic death in an automobile tion from the others.
accident in 2003, I assumed responsibility for completing This brings us to an important question: How are the
the 11th edition. separate organs and systems coordinated to maintain
For the 12th edition of the Textbook of Medical proper function of the entire body? Fortunately, our bod-
Physiology, I have the same goal as for previous editions— ies are endowed with a vast network of feedback con-
to explain, in language easily understood by students, how trols that achieve the necessary balances without which
the different cells, tissues, and organs of the human body we would be unable to live. Physiologists call this high
work together to maintain life. level of internal bodily control homeostasis. In disease
This task has been challenging and fun because our states, functional balances are often seriously disturbed
rapidly increasing knowledge of physiology continues to and homeostasis is impaired. When even a single distur-
unravel new mysteries of body functions. Advances in bance reaches a limit, the whole body can no longer live.
molecular and cellular physiology have made it possi- One of the goals of this text, therefore, is to emphasize the
ble to explain many physiology principles in the termi- effectiveness and beauty of the body’s homeostasis mech-
nology of molecular and physical sciences rather than anisms as well as to present their abnormal functions in
in merely a series of separate and unexplained biological disease.
phenomena. Another objective is to be as accurate as possible.
The Textbook of Medical Physiology, however, is not Suggestions and critiques from many students, physi-
a reference book that attempts to provide a compen- ologists, and clinicians throughout the world have been
dium of the most recent advances in physiology. This is sought and then used to check factual accuracy as well as
a book that continues the tradition of being written for balance in the text. Even so, because of the likelihood of
students. It focuses on the basic principles of physiol- error in sorting through many thousands of bits of infor-
ogy needed to begin a career in the health care profes- mation, I wish to issue a further request to all readers to
sions, such as medicine, dentistry and nursing, as well send along notations of error or inaccuracy. Physiologists
as graduate studies in the biological and health sciences. understand the importance of feedback for proper func-
It should also be useful to physicians and health care tion of the human body; so, too, is feedback important for
professionals who wish to review the basic principles progressive improvement of a textbook of physiology. To
needed for understanding the pathophysiology of the many persons who have already helped, I express sin-
human disease. cere thanks.
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Preface
A brief explanation is needed about several features of needed for immediate discussion but that most students
the 12th edition. Although many of the chapters have been will learn in more detail in other courses; second, physi-
revised to include new principles of physiology, the text ologic information of special importance to certain fields
length has been closely monitored to limit the book size of clinical medicine; and, third, information that will be of
so that it can be used effectively in physiology courses for value to those students who may wish to study particular
medical students and health care professionals. Many of the physiologic mechanisms more deeply.
figures have also been redrawn and are in full color. New ref- I wish to express sincere thanks to many persons who
erences have been chosen primarily for their presentation have helped to prepare this book, including my colleagues
of physiologic principles, for the quality of their own refer- in the Department of Physiology and Biophysics at the
ences, and for their easy accessibility. The selected biblio University of Mississippi Medical Center who provided
graphy at the end of the chapters lists papers mainly from valuable suggestions. The members of our faculty and a
recently published scientific journals that can be freely brief description of the research and educational activi-
accessed from the PubMed internet site at http://www. ties of the department can be found at the web site: http://
ncbi.nlm.nih.gov/sites/entrez/. Use of these references, as physiology.umc.edu/. I am also grateful to Stephanie
well as cross-references from them, can give the student Lucas and Courtney Horton Graham for their excellent
almost complete coverage of the entire field of physiology. secretarial services, to Michael Schenk and Walter (Kyle)
The effort to be as concise as possible has, unfortunately, Cunningham for their expert artwork, and to William
necessitated a more simplified and dogmatic presentation Schmitt, Rebecca Gruliow, Frank Morales, and the entire
of many physiologic principles than I normally would have Elsevier Saunders team for continued editorial and
desired. However, the bibliography can be used to learn production excellence.
more about the controversies and unanswered questions Finally, I owe an enormous debt to Arthur Guyton
that remain in understanding the complex functions of the for the great privilege of contributing to the Textbook of
human body in health and disease. Medical Physiology, for an exciting career in physiology,
Another feature is that the print is set in two sizes. The for his friendship, and for the inspiration that he provided
material in large print constitutes the fundamental physi- to all who knew him.
ologic information that students will require in virtually
all of their medical activities and studies. John E. Hall
The material in small print is of several different kinds:
first, anatomic, chemical, and other information that is
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Contents
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Contents
CHAPTER 6 CHAPTER 11
Contraction of Skeletal Muscle 71 The Normal Electrocardiogram 121
Physiologic Anatomy of Skeletal Muscle 71 Characteristics of the Normal
General Mechanism of Muscle Contraction 73 Electrocardiogram 121
Molecular Mechanism of Muscle Contraction 74 Methods for Recording Electrocardiograms 123
Energetics of Muscle Contraction 78 Flow of Current Around the Heart
Characteristics of Whole Muscle during the Cardiac Cycle 123
Contraction 79 Electrocardiographic Leads 124
CHAPTER 7 CHAPTER 12
Excitation of Skeletal Muscle: Electrocardiographic Interpretation of
Neuromuscular Transmission and Cardiac Muscle and Coronary Blood Flow
Excitation-Contraction Coupling 83 Abnormalities: Vectorial Analysis 129
Transmission of Impulses from Nerve Endings Principles of Vectorial Analysis of
to Skeletal Muscle Fibers: The Neuromuscular Electrocardiograms 129
Junction 83 Vectorial Analysis of the Normal
Molecular Biology of Acetylcholine Formation Electrocardiogram 131
and Release 86 Mean Electrical Axis of the Ventricular
Drugs That Enhance or Block Transmission QRS—and Its Significance 134
at the Neuromuscular Junction 86 Conditions That Cause Abnormal Voltages
Myasthenia Gravis Causes Muscle Paralysis 86 of the QRS Complex 137
Muscle Action Potential 87 Prolonged and Bizarre Patterns of the QRS
Excitation-Contraction Coupling 88 Complex 137
Current of Injury 138
CHAPTER 8
Abnormalities in the T Wave 141
Excitation and Contraction of Smooth Muscle 91
Contraction of Smooth Muscle 91 CHAPTER 13
Nervous and Hormonal Control of Smooth Cardiac Arrhythmias and Their
Muscle Contraction 94 Electrocardiographic Interpretation 143
Abnormal Sinus Rhythms 143
UNIT III Abnormal Rhythms That Result from Block
The Heart of Heart Signals Within the Intracardiac
Conduction Pathways 144
CHAPTER 9 Premature Contractions 146
Cardiac Muscle; The Heart as a Pump and Paroxysmal Tachycardia 148
Function of the Heart Valves 101 Ventricular Fibrillation 149
Physiology of Cardiac Muscle 101 Atrial Fibrillation 151
Cardiac Cycle 104 Atrial Flutter 152
Relationship of the Heart Sounds to Heart Cardiac Arrest 153
Pumping 107
Work Output of the Heart 107 UNIT IV
Chemical Energy Required for Cardiac Contraction:
The Circulation
Oxygen Utilization by the Heart 109
Regulation of Heart Pumping 110
CHAPTER 14
CHAPTER 10 Overview of the Circulation; Biophysics of
Rhythmical Excitation of the Heart 115 Pressure, Flow, and Resistance 157
Specialized Excitatory and Conductive System Physical Characteristics of the Circulation 157
of the Heart 115 Basic Principles of Circulatory Function 158
Control of Excitation and Conduction in the Interrelationships of Pressure, Flow, and
Heart 118 Resistance 159
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CHAPTER 15 CHAPTER 20
Vascular Distensibility and Functions of the Cardiac Output, Venous Return,
Arterial and Venous Systems 167 and Their Regulation 229
Vascular Distensibility 167 Normal Values for Cardiac Output at Rest
Arterial Pressure Pulsations 168 and During Activity 229
Veins and Their Functions 171 Control of Cardiac Output by Venous
Return—Role of the Frank-Starling Mechanism
CHAPTER 16 of the Heart 229
The Microcirculation and Lymphatic Pathologically High or Low Cardiac Outputs 232
System: Capillary Fluid Exchange, Methods for Measuring Cardiac
Interstitial Fluid, and Lymph Flow 177 Output 240
Structure of the Microcirculation
and Capillary System 177 CHAPTER 21
Flow of Blood in the Capillaries— Muscle Blood Flow and Cardiac Output
Vasomotion 178 During Exercise; the Coronary Circulation
and Ischemic Heart Disease 243
Exchange of Water, Nutrients, and Other
Substances Between the Blood and Blood Flow Regulation in Skeletal Muscle
Interstitial Fluid 179 at Rest and During Exercise 243
Interstitium and Interstitial Fluid 180 Coronary Circulation 246
Fluid Filtration Across Capillaries Is CHAPTER 22
Determined by Hydrostatic and Colloid
Osmotic Pressures, as Well as Capillary Cardiac Failure 255
Filtration Coefficient 181 Circulatory Dynamics in Cardiac Failure 255
Lymphatic System 186 Unilateral Left Heart Failure 259
Low-Output Cardiac Failure—
CHAPTER 17 Cardiogenic Shock 259
Local and Humoral Control of Tissue Edema in Patients with Cardiac Failure 259
Blood Flow 191
Cardiac Reserve 261
Local Control of Blood Flow in Response to
Tissue Needs 191 CHAPTER 23
Mechanisms of Blood Flow Control 191 Heart Valves and Heart Sounds;
Humoral Control of the Circulation 199 Valvular and Congenital Heart
Defects 265
CHAPTER 18 Heart Sounds 265
Nervous Regulation of the Circulation, Abnormal Circulatory Dynamics in Valvular
and Rapid Control of Arterial Pressure 201 Heart Disease 268
Nervous Regulation of the Circulation 201 Abnormal Circulatory Dynamics
Role of the Nervous System in Rapid in Congenital Heart Defects 269
Control of Arterial Pressure 204 Use of Extracorporeal Circulation During
Special Features of Nervous Control Cardiac Surgery 271
of Arterial Pressure 209 Hypertrophy of the Heart in Valvular
CHAPTER 19 and Congenital Heart Disease 272
Role of the Kidneys in Long-Term Control of CHAPTER 24
Arterial Pressure and in Hypertension: The
Circulatory Shock and Its Treatment 273
Integrated System for Arterial Pressure
Regulation 213 Physiologic Causes of Shock 273
Renal–Body Fluid System for Arterial Shock Caused by Hypovolemia—
Pressure Control 213 Hemorrhagic Shock 274
The Renin-Angiotensin System: Its Role Neurogenic Shock—Increased Vascular
in Arterial Pressure Control 220 Capacity 279
Summary of the Integrated, Multifaceted Anaphylactic Shock and Histamine Shock 280
System for Arterial Pressure Regulation 226 Septic Shock 280
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CHAPTER 36 CHAPTER 40
Hemostasis and Blood Coagulation 451 Transport of Oxygen and Carbon Dioxide in
Events in Hemostasis 451 Blood and Tissue Fluids 495
Vascular Constriction 451 Transport of Oxygen from the Lungs to the
Mechanism of Blood Coagulation 453 Body Tissues 495
Conditions That Cause Excessive Bleeding in Transport of Carbon Dioxide in the Blood 502
Humans 457 Respiratory Exchange Ratio 504
Thromboembolic Conditions in the CHAPTER 41
Human Being 459 Regulation of Respiration 505
Anticoagulants for Clinical Use 459 Respiratory Center 505
Blood Coagulation Tests 460 Chemical Control of Respiration 507
Peripheral Chemoreceptor System for Control
UNIT VII of Respiratory Activity—Role of Oxygen in
Respiration Respiratory Control 508
Regulation of Respiration During Exercise 510
CHAPTER 37 Other Factors That Affect Respiration 512
Pulmonary Ventilation 465
CHAPTER 42
Mechanics of Pulmonary Ventilation 465
Respiratory Insufficiency—Pathophysiology,
Pulmonary Volumes and Capacities 469 Diagnosis, Oxygen Therapy 515
Minute Respiratory Volume Equals Respiratory Useful Methods for Studying Respiratory
Rate Times Tidal Volume 471 Abnormalities 515
Alveolar Ventilation 471 Pathophysiology of Specific Pulmonary
Functions of the Respiratory Passageways 472 Abnormalities 517
Hypoxia and Oxygen Therapy 520
CHAPTER 38 Hypercapnia—Excess Carbon Dioxide in the
Pulmonary Circulation, Pulmonary Edema, Body Fluids 522
Pleural Fluid 477 Artificial Respiration 522
Physiologic Anatomy of the Pulmonary
Circulatory System 477 UNIT VIII
Pressures in the Pulmonary System 477 Aviation, Space, and Deep-Sea Diving
Blood Volume of the Lungs 478 Physiology
Blood Flow Through the Lungs and Its
Distribution 479 CHAPTER 43
Effect of Hydrostatic Pressure Gradients in Aviation, High-Altitude, and
the Lungs on Regional Pulmonary Blood Flow 479 Space Physiology 527
Pulmonary Capillary Dynamics 481 Effects of Low Oxygen Pressure on the Body 527
Fluid in the Pleural Cavity 483 Effects of Acceleratory Forces on the Body in
Aviation and Space Physiology 531
CHAPTER 39 “Artificial Climate” in the Sealed Spacecraft 533
Physical Principles of Gas Exchange; Weightlessness in Space 533
Diffusion of Oxygen and Carbon Dioxide
Through the Respiratory Membrane 485 CHAPTER 44
Physics of Gas Diffusion and Gas Physiology of Deep-Sea Diving and
Partial Pressures 485 Other Hyperbaric Conditions 535
Compositions of Alveolar Air and Atmospheric Effect of High Partial Pressures of Individual
Air Are Different 487 Gases on the Body 535
Diffusion of Gases Through the Respiratory Scuba (Self-Contained Underwater Breathing
Membrane 489 Apparatus) Diving 539
Effect of the Ventilation-Perfusion Ratio on Special Physiologic Problems in Submarines 540
Alveolar Gas Concentration 492 Hyperbaric Oxygen Therapy 540
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CHAPTER 71 CHAPTER 75
Dietary Balances; Regulation of Feeding; Pituitary Hormones and Their Control by the
Obesity and Starvation; Vitamins and Hypothalamus 895
Minerals 843 Pituitary Gland and Its Relation to the
Energy Intake and Output Are Balanced Under Hypothalamus 895
Steady-State Conditions 843 Hypothalamus Controls Pituitary Secretion 897
Dietary Balances 843 Physiological Functions of Growth Hormone 898
Regulation of Food Intake and Energy Posterior Pituitary Gland and Its Relation to
Storage 845 the Hypothalamus 904
Obesity 850
CHAPTER 76
Inanition, Anorexia, and Cachexia 851
Thyroid Metabolic Hormones 907
Starvation 852
Synthesis and Secretion of the Thyroid
Vitamins 852
Metabolic Hormones 907
Mineral Metabolism 855
Physiological Functions of the Thyroid
CHAPTER 72 Hormones 910
Energetics and Metabolic Rate 859 Regulation of Thyroid Hormone Secretion 914
Adenosine Triphosphate (ATP) Functions as Diseases of the Thyroid 916
an “Energy Currency” in Metabolism 859 CHAPTER 77
Control of Energy Release in the Cell 861 Adrenocortical Hormones 921
Metabolic Rate 862 Synthesis and Secretion of Adrenocortical
Energy Metabolism—Factors That Influence Hormones 921
Energy Output 863 Functions of the Mineralocorticoids—
CHAPTER 73 Aldosterone 924
Body Temperature Regulation, Functions of the Glucocorticoids 928
and Fever 867 Adrenal Androgens 934
Normal Body Temperatures 867 Abnormalities of Adrenocortical Secretion 934
Body Temperature Is Controlled by CHAPTER 78
Balancing Heat Production and
Heat Loss 867 Insulin, Glucagon, and Diabetes Mellitus 939
Regulation of Body Temperature— Insulin and Its Metabolic Effects 939
Role of the Hypothalamus 871 Glucagon and Its Functions 947
Abnormalities of Body Temperature Somatostatin Inhibits Glucagon and Insulin
Regulation 875 Secretion 949
Summary of Blood Glucose Regulation 949
Diabetes Mellitus 950
UNIT XIV
CHAPTER 79
Endocrinology and Reproduction Parathyroid Hormone, Calcitonin, Calcium
and Phosphate Metabolism, Vitamin D, Bone,
CHAPTER 74 and Teeth 955
Introduction to Endocrinology 881 Overview of Calcium and
Coordination of Body Functions by Chemical Phosphate Regulation in the Extracellular
Messengers 881 Fluid and Plasma 955
Chemical Structure and Synthesis of Bone and Its Relation to Extracellular Calcium
Hormones 881 and Phosphate 957
Hormone Secretion, Transport, and Clearance Vitamin D 960
from the Blood 884 Parathyroid Hormone 962
Mechanisms of Action of Hormones 886 Calcitonin 966
Measurement of Hormone Concentrations Summary of Control of Calcium Ion
in the Blood 891 Concentration 966
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