2. T E X T B O O K of Medical Physiology E L E V E N T H E D I
T I O N Arthur C. Guyton, M.D. Professor Emeritus Department of
Physiology and Biophysics University of Mississippi Medical Center
Jackson, Mississippi Deceased John E. Hall, Ph.D. Professor and
Chairman Department of Physiology and Biophysics University of
Mississippi Medical Center Jackson, Mississippi
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medical physiology / Arthur C. Guyton, John E. Hall.11th ed. p. ;
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0-7216-0240-1 1. Human physiology. 2. Physiology, Pathological. I.
Title: Medical physiology. II. Hall, John E. (John Edward) III.
Title. [DNLM: 1. Physiological Processes. QT 104 G992t 2006]
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4. 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
5. Arthur C. Guyton, M.D. 19192003
6. I N M E M O R I A M The sudden loss of Dr. Arthur C. Guyton
in an automobile accident on April 3, 2003, stunned and saddened
all who were privileged to know him. Arthur Guyton was a giant in
the elds of physiology and medicine, a leader among leaders, a
master teacher, and an inspiring role model throughout the world.
Arthur Clifton Guyton was born in Oxford, Mississippi, to Dr. Billy
S. Guyton, a highly respected eye, ear, nose, and throat
specialist, who later became Dean of the University of Mississippi
Medical School, and Kate Small- wood Guyton, a mathematics and
physics teacher who had been a missionary in China before marriage.
During his formative years,Arthur enjoyed watching his father work
at the Guyton Clinic, playing chess and swapping stories with
William Faulkner, and building sailboats (one of which he later
sold to Faulkner). He also built countless mechanical and
electrical devices, which he continued to do throughout his life.
His brilliance shone early as he graduated top in his class at the
University of Mississippi. He later distinguished himself at
Harvard Medical School and began his postgraduate surgical training
at Massachusetts General Hospital. His medical training was
interrupted twiceonce to serve in the Navy during World War II and
again in 1946 when he was stricken with poliomyelitis during his
nal year of residency training. Suffering paralysis in his right
leg, left arm, and both shoulders, he spent nine months in Warm
Springs, Georgia, recuper- ating and applying his inventive mind to
building the rst motorized wheelchair controlled by a joy stick, a
motorized hoist for lifting patients, special leg braces, and other
devices to aid the handicapped. For those inventions he received a
Presidential Citation. He returned to Oxford where he devoted
himself to teaching and research at the University of Mississippi
School of Medicine and was named Chair of the Department of
Physiology in 1948. In 1951 he was named one of the ten out-
standing men in the nation. When the University of Mississippi
moved its Medical School to Jackson in 1955, he rapidly developed
one of the worlds premier cardiovascular research programs. His
remarkable life as a scientist, author, and devoted father is
detailed in a biography published on the occasion of his retirement
in 1989.1 A Great Physiologist. Arthur Guytons research
contributions, which include more than 600 papers and 40 books, are
legendary and place him among the greatest physiologists in
history. His research covered virtually all areas of car-
diovascular regulation and led to many seminal concepts that are
now an inte- gral part of our understanding of cardiovascular
disorders, such as hypertension, heart failure, and edema. It is
difcult to discuss cardiovascular physiology without including his
concepts of cardiac output and venous return, negative interstitial
uid pressure and regulation of tissue uid volume and edema,
regulation of tissue blood ow and whole body blood ow
autoregulation, renal-pressure natriuresis, and long-term blood
pressure regulation. Indeed, his concepts of cardiovascular
regulation are found in virtually every major text- book of
physiology.They have become so familiar that their origin is
sometimes forgotten. One of Dr. Guytons most important scientic
legacies was his application of principles of engineering and
systems analysis to cardiovascular regulation. He used mathematical
and graphical methods to quantify various aspects of circu- latory
function before computers were widely available. He built analog
com- puters and pioneered the application of large-scale systems
analysis to modeling the cardiovascular system before the advent of
digital computers. As digital computers became available, his
cardiovascular models expanded dramatically to include the kidneys
and body uids, hormones, and the autonomic nervous system, as well
as cardiac and circulatory functions.2 He also provided the rst
comprehensive systems analysis of blood pressure regulation. This
unique approach to physiological research preceded the emergence of
biomedical vii
7. viii In Memoriam engineeringa eld that he helped to
establish and to promote in physiology, leading the discipline into
a quantitative rather than a descriptive science. It is a tribute
to Arthur Guytons genius that his concepts of cardiovascular
regulation often seemed heretical when they were rst presented, yet
stimu- lated investigators throughout the world to test them
experimentally.They are now widely accepted. In fact, many of his
concepts of cardiovascular regulation are integral components of
what is now taught in most medical physiology courses. They
continue to be the foundation for generations of cardiovascular
physiologists. Dr. Guyton received more than 80 major honors from
diverse scientic and civic organizations and uni- versities
throughout the world.A few of these that are especially relevant to
cardiovascular research include the Wiggers Award of the American
Physiological Society, the Ciba Award from the Council for High
Blood Pressure Research, The William Harvey Award from the American
Society of Hypertension, the Research Achievement Award of the
American Heart Association, and the Merck Sharp & Dohme Award
of the International Society of Hypertension. It was appropriate
that in 1978 he was invited by the Royal College of Physicians in
London to deliver a special lecture honoring the 400th anniversary
of the birth of William Harvey, who discovered the circulation of
the blood. Dr. Guytons love of physiology was beautifully
articulated in his presidents address to the American Physiological
Society in 1975,3 appropriately entitled Physiology, a Beauty and a
Philosophy. Let me quote just one sentence from his address: What
other person, whether he be a theologian, a jurist, a doctor of
medi- cine, a physicist, or whatever, knows more than you, a
physiologist, about life? For physiology is indeed an explanation
of life. What other subject matter is more fascinating, more
exciting, more beautiful than the subject of life? A Master
Teacher. Although Dr. Guytons research accomplishments are
legendary, his contributions as an educator have probably had an
even greater impact. He and his wonderful wife Ruth raised ten
children, all of whom became outstanding physiciansa remarkable
educational achievement. Eight of the Guyton children graduated
from Harvard Medical School, one from Duke Medical School, and one
from The University of Miami Medical School after receiv- ing a
Ph.D. from Harvard. An article published in Readers Digest in 1982
highlighted their extraordinary family life.4 The success of the
Guyton children did not occur by chance. Dr. Guytons philosophy of
education was to learn by doing. The children participated in
count- less family projects that included the design and
construction of their home and its heating system, the swimming
pool, tennis court, sailboats, go-carts and electrical cars,
household gadgets, and electronic instruments for their Oxford
Instruments Company. Television programs such as Good Morning
America and 20/20 described the remarkable home environ- ment that
Arthur and Ruth Guyton created to raise their family. His devotion
to family is beautifully expressed in the dedication of his
Textbook of Medical Physiology5 : To My father for his
uncompromising principles that guided my life My mother for leading
her children into intellectual pursuits My wife for her magnicent
devotion to her family My children for making everything worthwhile
Dr. Guyton was a master teacher at the University of Mississippi
for over 50 years. Even though he was always busy with service
responsibilities, research, writing, and teaching, he was never too
busy to talk with a student who was having difculty. He would never
accept an invitation to give a prestigious lecture if it conicted
with his teaching schedule. His contributions to education are also
far reach- ing through generations of physiology graduate students
and postdoctoral fellows. He trained over 150 scientists, at least
29 of whom became chairs of their own departments and six of whom
became pres- idents of the American Physiological Society. He gave
students condence in their abilities and emphasized his belief that
People who are really successful in the research world are
self-taught. He insisted that his trainees integrate their
experimental ndings into a broad conceptual framework that included
other interacting systems. This approach usually led them to
develop a quantitative analysis and a better understanding of the
particular physiological systems that they were studying. No one
has been more pro- lic in training leaders of physiology than
Arthur Guyton. Dr. Guytons Textbook of Medical Physiology, rst
published in 1956, quickly became the best-selling medical
physiology textbook in the world. He had a gift for communicating
complex ideas in a clear and interesting manner that made studying
physiology fun. He wrote the book to teach his students, not to
impress his professional colleagues. Its popularity with stu- dents
has made it the most widely used physiology textbook in history.
This accomplishment alone was enough to ensure his legacy. The
Textbook of Medical Physiology began as lecture notes in the early
1950s when Dr. Guyton was teaching the entire physiology course for
medical stu- dents at the University of Mississippi. He discovered
that the students were having difculty with the text- books that
were available and began distributing copies of his lecture notes.
In describing his experi- ence, Dr. Guyton stated that Many
textbooks of medical physiology had become discursive, written pri-
marily by teachers of physiology for other teachers of physiology,
and written in language understood by other teachers but not easily
understood by the basic student of medical physiology.6 Through his
Textbook of Medical Physiology, which is translated into 13
languages, he has probably done
8. In Memoriam ix more to teach physiology to the world than
any other individual in history. Unlike most major textbooks, which
often have 20 or more authors, the rst eight editions were written
entirely by Dr. Guytona feat that is unprecedented for any major
medical textbook. For his many contributions to medical education,
Dr. Guyton received the 1996 Abraham Flexner Award from the
Association of American Medical Colleges (AAMC). According to the
AAMC, Arthur Guyton . . . for the past 50 years has made an
unparalleled impact on medical education. He is also honored each
year by The American Physiological Society through the Arthur C.
Guyton Teaching Award. An Inspiring Role Model. Dr. Guytons
accomplish- ments extended far beyond science,medicine,and edu-
cation. He was an inspiring role model for life as well as for
science. No one was more inspirational or inu- ential on my
scientic career than Dr. Guyton. He taught his students much more
than physiology he taught us life, not so much by what he said but
by his unspoken courage and dedication to the highest standards. He
had a special ability to motivate people through his indomitable
spirit. Although he was severely chal- lenged by polio, those of us
who worked with him never thought of him as being handicapped. We
were too busy trying to keep up with him! His brilliant mind, his
indefatigable devotion to science, education, and family, and his
spirit captivated students and trainees, professional colleagues,
politicians, business leaders, and virtually everyone who knew him.
He would not succumb to the effects of polio. His courage
challenged and inspired us. He expected the best and somehow
brought out the very best in people. We celebrate the magnicent
life of Arthur Guyton, recognizing that we owe him an enormous
debt. He gave us an imaginative and innovative approach to research
and many new scientic concepts. He gave countless students
throughout the world a means of understanding physiology and he
gave many of us exciting research careers. Most of all, he inspired
us with his devotion to education, his unique ability to bring out
the best in those around him, his warm and generous spirit, and his
courage. We will miss him tremendously, but he will remain in our
memories as a shining example of the very best in humanity.Arthur
Guyton was a real hero to the world, and his legacy is everlasting.
References 1. Brinson C, Quinn J: Arthur C. GuytonHis Life, His
Family, His Achievements. Jackson, MS, Hederman Brothers Press,
1989. 2. Guyton AC, Coleman TG, Granger HJ: Circulation: overall
regulation. Ann Rev Physiol 34:1346, 1972. 3. Guyton AC:
Past-Presidents Address. Physiology, a Beauty and a Philosophy. The
Physiologist 8:495501, 1975. 4. Bode R:A Doctor Whos Dad to Seven
DoctorsSo Far! Readers Digest, December, 1982, pp. 141145. 5.
Guyton AC: Textbook of Medical Physiology. Philadel- phia,
Saunders, 1956. 6. Guyton AC: An authors philosophy of physiology
text- book writing. Adv Physiol Ed 19: s1s5, 1998. John E. Hall
Jackson, Mississippi
9. P R E F A C E The rst edition of the Textbook of Medical
Phys- iology was written by Arthur C. Guyton almost 50 years ago.
Unlike many major medical textbooks, which often have 20 or more
authors, the rst eight editions of the Textbook of Medical Physi-
ology were written entirely by Dr. Guyton with each new edition
arriving on schedule for nearly 40 years. Over the years, Dr.
Guytons textbook became widely used throughout the world and was
translated into 13 languages. A major reason for the books
unprecedented success was his uncanny ability to explain complex
physiologic principles in language easily understood by stu- dents.
His main goal with each edition was to instruct students in
physiology, not to impress his professional colleagues. His writing
style always maintained the tone of a teacher talking to his
students. I had the privilege of working closely with Dr. Guyton
for almost 30 years and the honor of helping him with the 9th and
10th editions. For the 11th edition, I have the same goal as in
previous editionsto explain, in language easily understood by
students, how the different cells, tissues, and organs of the human
body work together to maintain life. This task has been challenging
and exciting because our rapidly increasing knowledge of physiology
continues to unravel new mysteries of body functions. Many new
techniques for learning about molecular and cellular physiology
have been developed. We can present more and more the physiology
principles in the terminology of molecular and physical sciences
rather than in merely a series of separate and unexplained bio-
logical phenomena.This change is welcomed, but it also makes
revision of each chapter a necessity. In this edition, I have
attempted to maintain the same unied organization of the text that
has been useful to students in the past and to ensure that the book
is comprehensive enough that students will wish to use it in later
life as a basis for their professional careers. I hope that this
textbook conveys the majesty of the human body and its many
functions and that it stimulates students to study physiology
throughout their careers. Physiology is the link between the basic
sciences and medicine. The great beauty of physiology is that it
integrates the individual functions of all the bodys different
cells, tissues, and organs into a functional whole, the human body.
Indeed, the human body is much more than the sum of its parts, and
life relies upon this total func- tion, not just on the function of
individual body parts in isolation from the others. This brings us
to an important question: How are the separate organs and systems
coordinated to maintain proper function of the entire body? Fortu-
nately, our bodies are endowed with a vast network of feedback
controls that achieve the necessary balances without which we would
not be able to live. Physiologists call this high level of internal
bodily control homeostasis. In disease states, functional balances
are often seriously disturbed and homeosta- sis is impaired. And,
when even a single disturbance reaches a limit, the whole body can
no longer live. One of the goals of this text, therefore, is to
emphasize the effectiveness and beauty of the bodys homeostasis
mechanisms as well as to present their abnormal function in
disease. Another objective is to be as accurate as possible.
Suggestions and critiques from many physiologists, students, and
clinicians throughout the world have been sought and then used to
check factual accuracy as well as balance in the text. Even so,
because of the likelihood of error in sorting through many thou-
sands of bits of information, I wish to issue still a further
request to all readers to send along notations of error or
inaccuracy. Physiologists understand the importance of feedback for
proper function of the human body; so, too, is feed- back important
for progressive improvement of a textbook of physiology. To the
many persons who have already helped, I send sincere thanks.
xi
10. xii Preface A brief explanation is needed about several
features of the 11th edition. Although many of the chapters have
been revised to include new principles of physi- ology, the text
length has been closely monitored to limit the book size so that it
can be used effec- tively in physiology courses for medical
students and health care professionals. Many of the gures have also
been redrawn and are now in full color. New references have been
chosen primarily for their pres- entation of physiologic
principles, for the quality of their own references, and for their
easy accessibility. Most of the selected references are from
recently published scientic journals that can be freely accessed
from the PubMed internet site at http://
www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed. Use of these
references, as well as cross-references from them, can give the
student almost complete cov- erage of the entire eld of physiology.
Another feature is that the print is set in two sizes. The material
in small print is of several different kinds: rst, anatomical,
chemical, and other information that is needed for immediate
discussion but that most stu- dents will learn in more detail in
other courses; second, physiologic information of special
importance to certain elds of clinical medicine; and, third,
informa- tion that will be of value to those students who may wish
to study particular physiologic mechanisms more deeply. The
material in large print constitutes the funda- mental physiologic
information that students will require in virtually all their
medical activities and studies. I wish to express my thanks to many
other persons who have helped in preparing this book, including my
colleagues in the Department of Physiology & Biophysics at the
University of Mississippi Medical Center who provided valuable
suggestions. I am also grateful to Ivadelle Osberg Heidke, Gerry
McAlpin, and Stephanie Lucas for their excellent secretarial
services, and to William Schmitt, Rebecca Gruliow, Mary Anne
Folcher, and the rest of the staff of Elsevier Saunders for
continued editorial and produc- tion excellence. Finally, I owe an
enormous debt to Arthur Guyton for an exciting career in
physiology, for his friendship, for the great privilege of
contributing to the Textbook of Medical Physiology, and for the
inspiration that he provided to all who knew him. John E. Hall
Jackson, Mississippi
11. T A B L E O F C O N T E N T S U N I T I Introduction to
Physiology: The Cell and General Physiology C H A P T E R 1
Functional Organization of the Human Body and Control of the
Internal Environment 3 Cells as the Living Units of the Body 3
Extracellular FluidThe Internal Environment 3 Homeostatic
Mechanisms of the Major Functional Systems 4 Homeostasis 4
Extracellular Fluid Transport and Mixing SystemThe Blood
Circulatory System 4 Origin of Nutrients in the Extracellular Fluid
5 Removal of Metabolic End Products 5 Regulation of Body Functions
5 Reproduction 6 Control Systems of the Body 6 Examples of Control
Mechanisms 6 Characteristics of Control Systems 7
SummaryAutomaticity of the Body 9 C H A P T E R 2 The Cell and Its
Functions 11 Organization of the Cell 11 Physical Structure of the
Cell 12 Membranous Structures of the Cell 12 Cytoplasm and Its
Organelles 14 Nucleus 17 Nuclear Membrane 17 Nucleoli and Formation
of Ribosomes 18 Comparison of the Animal Cell with Precellular
Forms of Life 18 Functional Systems of the Cell 19 Ingestion by the
CellEndocytosis 19 Digestion of Pinocytotic and Phagocytic Foreign
Substances Inside the Cell Function of the Lysosomes 20 Synthesis
and Formation of Cellular Structures by Endoplasmic Reticulum and
Golgi Apparatus 20 Extraction of Energy from Nutrients Function of
the Mitochondria 22 Locomotion of Cells 24 Ameboid Movement 24
Cilia and Ciliary Movement 24 C H A P T E R 3 Genetic Control of
Protein Synthesis, Cell Function, and Cell Reproduction 27 Genes in
the Cell Nucleus 27 Genetic Code 29 xiii The DNA Code in the Cell
Nucleus Is Transferred to an RNA Code in the Cell CytoplasmThe
Process of Transcription 30 Synthesis of RNA 30 Assembly of the RNA
Chain from Activated Nucleotides Using the DNA Strand as a
TemplateThe Process of Transcription 31 Messenger RNAThe Codons 31
Transfer RNAThe Anticodons 32 Ribosomal RNA 33 Formation of
Proteins on the Ribosomes The Process of Translation 33 Synthesis
of Other Substances in the Cell 35 Control of Gene Function and
Biochemical Activity in Cells 35 Genetic Regulation 35 Control of
Intracellular Function by Enzyme Regulation 36 The DNA-Genetic
System Also Controls Cell Reproduction 37 Cell Reproduction Begins
with Replication of DNA 37 Chromosomes and Their Replication 38
Cell Mitosis 38 Control of Cell Growth and Cell Reproduction 39
Cell Differentiation 40 ApoptosisProgrammed Cell Death 40 Cancer 40
U N I T I I Membrane Physiology, Nerve, and Muscle C H A P T E R 4
Transport of Substances Through the Cell Membrane 45 The Lipid
Barrier of the Cell Membrane, and Cell Membrane Transport Proteins
45 Diffusion 46 Diffusion Through the Cell Membrane 46 Diffusion
Through Protein Channels, and Gating of These Channels 47
Facilitated Diffusion 49 Factors That Affect Net Rate of Diffusion
50 Osmosis Across Selectively Permeable MembranesNet Diffusion of
Water 51 Active Transport of Substances Through Membranes 52
Primary Active Transport 53 Secondary Active TransportCo-Transport
and Counter-Transport 54 Active Transport Through Cellular Sheets
55
12. xiv Table of Contents C H A P T E R 5 Membrane Potentials
and Action Potentials 57 Basic Physics of Membrane Potentials 57
Membrane Potentials Caused by Diffusion 57 Measuring the Membrane
Potential 58 Resting Membrane Potential of Nerves 59 Origin of the
Normal Resting Membrane Potential 60 Nerve Action Potential 61
Voltage-Gated Sodium and Potassium Channels 62 Summary of the
Events That Cause the Action Potential 64 Roles of Other Ions
During the Action Potential 64 Initiation of the Action Potential
65 Propagation of the Action Potential 65 Re-establishing Sodium
and Potassium Ionic Gradients After Action Potentials Are
CompletedImportance of Energy Metabolism 66 Plateau in Some Action
Potentials 66 Rhythmicity of Some Excitable Tissues Repetitive
Discharge 67 Special Characteristics of Signal Transmission in
Nerve Trunks 68 ExcitationThe Process of Eliciting the Action
Potential 69 Refractory Period After an Action Potential 70
Recording Membrane Potentials and Action Potentials 70 Inhibition
of ExcitabilityStabilizers and Local Anesthetics 70 C H A P T E R 6
Contraction of Skeletal Muscle 72 Physiologic Anatomy of Skeletal
Muscle 72 Skeletal Muscle Fiber 72 General Mechanism of Muscle
Contraction 74 Molecular Mechanism of Muscle Contraction 74
Molecular Characteristics of the Contractile Filaments 75 Effect of
Amount of Actin and Myosin Filament Overlap on Tension Developed by
the Contracting Muscle 77 Relation of Velocity of Contraction to
Load 78 Energetics of Muscle Contraction 78 Work Output During
Muscle Contraction 78 Sources of Energy for Muscle Contraction 79
Characteristics of Whole Muscle Contraction 80 Mechanics of
Skeletal Muscle Contraction 81 Remodeling of Muscle to Match
Function 82 Rigor Mortis 83 C H A P T E R 7 Excitation of Skeletal
Muscle: Neuromuscular Transmission and Excitation-Contraction
Coupling 85 Transmission of Impulses from Nerve Endings to Skeletal
Muscle Fibers: The Neuromuscular Junction 85 Secretion of
Acetylcholine by the Nerve Terminals 85 Molecular Biology of
Acetyline Formation and Release 88 Drugs That Enhance or Block
Transmission at the Neuromuscular Junction 88 Myasthenia Gravis 89
Muscle Action Potential 89 Spread of the Action Potential to the
Interior of the Muscle Fiber by Way of Transverse Tubules 89
Excitation-Contraction Coupling 89 Transverse TubuleSarcoplasmic
Reticulum System 89 Release of Calcium Ions by the Sarcoplasmic
Reticulum 90 C H A P T E R 8 Contraction and Excitation of Smooth
Muscle 92 Contraction of Smooth Muscle 92 Types of Smooth Muscle 92
Contractile Mechanism in Smooth Muscle 93 Regulation of Contraction
by Calcium Ions 95 Nervous and Hormonal Control of Smooth Muscle
Contraction 95 Neuromuscular Junctions of Smooth Muscle 95 Membrane
Potentials and Action Potentials in Smooth Muscle 96 Effect of
Local Tissue Factors and Hormones to Cause Smooth Muscle
Contraction Without Action Potentials 98 Source of Calcium Ions
That Cause Contraction (1) Through the Cell Membrane and (2) from
the Sarcoplasmic Reticulum 99 U N I T I I I The Heart C H A P T E R
9 Heart Muscle; The Heart as a Pump and Function of the Heart
Valves 103 Physiology of Cardiac Muscle 103 Physiologic Anatomy of
Cardiac Muscle 103 Action Potentials in Cardiac Muscle 104 The
Cardiac Cycle 106 Diastole and Systole 106 Relationship of the
Electrocardiogram to the Cardiac Cycle 107 Function of the Atria as
Primer Pumps 107 Function of the Ventricles as Pumps 108
13. Table of Contents xv Function of the Valves 109 Aortic
Pressure Curve 109 Relationship of the Heart Sounds to Heart
Pumping 109 Work Output of the Heart 110 Graphical Analysis of
Ventricular Pumping 110 Chemical Energy Required for Cardiac
Contraction: Oxygen Utilization by the Heart 111 Regulation of
Heart Pumping 111 Intrinsic Regulation of Heart Pumping The
Frank-Starling Mechanism 111 Effect of Potassium and Calcium Ions
on Heart Function 113 Effect of Temperature on Heart Function 114
Increasing the Arterial Pressure Load (up to a Limit) Does Not
Decrease the Cardiac Output 114 C H A P T E R 1 0 Rhythmical
Excitation of the Heart 116 Specialized Excitatory and Conductive
System of the Heart 116 Sinus (Sinoatrial) Node 116 Internodal
Pathways and Transmission of the Cardiac Impulse Through the Atria
118 Atrioventricular Node, and Delay of Impulse Conduction from the
Atria to the Ventricles 118 Rapid Transmission in the Ventricular
Purkinje System 119 Transmission of the Cardiac Impulse in the
Ventricular Muscle 119 Summary of the Spread of the Cardiac Impulse
Through the Heart 120 Control of Excitation and Conduction in the
Heart 120 The Sinus Node as the Pacemaker of the Heart 120 Role of
the Purkinje System in Causing Synchronous Contraction of the
Ventricular Muscle 121 Control of Heart Rhythmicity and Impulse
Conduction by the Cardiac Nerves: The Sympathetic and
Parasympathetic Nerves 121 C H A P T E R 1 1 The Normal
Electrocardiogram 123 Characteristics of the Normal
Electrocardiogram 123 Depolarization Waves Versus Repolarization
Waves 123 Relationship of Atrial and Ventricular Contraction to the
Waves of the Electrocardiogram 125 Voltage and Time Calibration of
the Electrocardiogram 125 Methods for Recording Electrocardiograms
126 Pen Recorder 126 Flow of Current Around the Heart During the
Cardiac Cycle 126 Recording Electrical Potentials from a Partially
Depolarized Mass of Syncytial Cardiac Muscle 126 Flow of Electrical
Currents in the Chest Around the Heart 126 Electrocardiographic
Leads 127 Three Bipolar Limb Leads 127 Chest Leads (Precordial
Leads) 129 Augmented Unipolar Limb Leads 129 C H A P T E R 1 2
Electrocardiographic Interpretation of Cardiac Muscle and Coronary
Blood Flow Abnormalities: Vectorial Analysis 131 Principles of
Vectorial Analysis of Electrocardiograms 131 Use of Vectors to
Represent Electrical Potentials 131 Direction of a Vector Is
Denoted in Terms of Degrees 131 Axis for Each Standard Bipolar Lead
and Each Unipolar Limb Lead 132 Vectorial Analysis of Potentials
Recorded in Different Leads 133 Vectorial Analysis of the Normal
Electrocardiogram 134 Vectors That Occur at Successive Intervals
During Depolarization of the Ventricles The QRS Complex 134
Electrocardiogram During Repolarization The T Wave 134
Depolarization of the AtriaThe P Wave 136 Vectorcardiogram 136 Mean
Electrical Axis of the Ventricular QRSAnd Its Signicance 137
Determining the Electrical Axis from Standard Lead
Electrocardiograms 137 Abnormal Ventricular Conditions That Cause
Axis Deviation 138 Conditions That Cause Abnormal Voltages of the
QRS Complex 140 Increased Voltage in the Standard Bipolar Limb
Leads 140 Decreased Voltage of the Electrocardiogram 140 Prolonged
and Bizarre Patterns of the QRS Complex 141 Prolonged QRS Complex
as a Result of Cardiac Hypertrophy or Dilatation 141 Prolonged QRS
Complex Resulting from Purkinje System Blocks 141 Conditions That
Cause Bizarre QRS Complexes 141 Current of Injury 141 Effect of
Current of Injury on the QRS Complex 141 The J PointThe Zero
Reference Potential for Analyzing Current of Injury 142 Coronary
Ischemia as a Cause of Injury Potential 143 Abnormalities in the T
Wave 145 Effect of Slow Conduction of the Depolarization Wave on
the Characteristics of the T Wave 145 Shortened Depolarization in
Portions of the Ventricular Muscle as a Cause of T Wave
Abnormalities 145
14. xvi Table of Contents C H A P T E R 1 3 Cardiac Arrhythmias
and Their Electrocardiographic Interpretation 147 Abnormal Sinus
Rhythms 147 Tachycardia 147 Bradycardia 147 Sinus Arrhythmia 148
Abnormal Rhythms That Result from Block of Heart Signals Within the
Intracardiac Conduction Pathways 148 Sinoatrial Block 148
Atrioventricular Block 148 Incomplete Atrioventricular Heart Block
149 Incomplete Intraventricular Block Electrical Alternans 150
Premature Contractions 150 Premature Atrial Contractions 150 A-V
Nodal or A-V Bundle Premature Contractions 150 Premature
Ventricular Contractions 151 Paroxysmal Tachycardia 151 Atrial
Paroxysmal Tachycardia 152 Ventricular Paroxysmal Tachycardia 152
Ventricular Fibrillation 152 Phenomenon of Re-entryCircus Movements
as the Basis for Ventricular Fibrillation 153 Chain Reaction
Mechanism of Fibrillation 153 Electrocardiogram in Ventricular
Fibrillation 154 Electroshock Debrillation of the Ventricle 154
Hand Pumping of the Heart (Cardiopulmonary Resuscitation) as an Aid
to Debrillation 155 Atrial Fibrillation 155 Atrial Flutter 156
Cardiac Arrest 156 U N I T I V The Circulation C H A P T E R 1 4
Overview of the Circulation; Medical Physics of Pressure, Flow, and
Resistance 161 Physical Characteristics of the Circulation 161
Basic Theory of Circulatory Function 163 Interrelationships Among
Pressure, Flow, and Resistance 164 Blood Flow 164 Blood Pressure
166 Resistance to Blood Flow 167 Effects of Pressure on Vascular
Resistance and Tissue Blood Flow 170 C H A P T E R 1 5 Vascular
Distensibility and Functions of the Arterial and Venous Systems 171
Vascular Distensibility 171 Vascular Compliance (or Vascular
Capacitance) 171 Volume-Pressure Curves of the Arterial and Venous
Circulations 172 Arterial Pressure Pulsations 173 Transmission of
Pressure Pulses to the Peripheral Arteries 174 Clinical Methods for
Measuring Systolic and Diastolic Pressures 175 Veins and Their
Functions 176 Venous PressuresRight Atrial Pressure (Central Venous
Pressure) and Peripheral Venous Pressures 176 Blood Reservoir
Function of the Veins 179 C H A P T E R 1 6 The Microcirculation
and the Lymphatic System: Capillary Fluid Exchange, Interstitial
Fluid, and Lymph Flow 181 Structure of the Microcirculation and
Capillary System 181 Flow of Blood in the Capillaries Vasomotion
182 Average Function of the Capillary System 183 Exchange of Water,
Nutrients, and Other Substances Between the Blood and Interstitial
Fluid 183 Diffusion Through the Capillary Membrane 183 The
Interstitium and Interstitial Fluid 184 Fluid Filtration Across
Capillaries Is Determined by Hydrostatic and Colloid Osmotic
Pressures, and Capillary Filtration Coefcient 185 Capillary
Hydrostatic Pressure 186 Interstitial Fluid Hydrostatic Pressure
187 Plasma Colloid Osmotic Pressure 188 Interstitial Fluid Colloid
Osmotic Pressure 188 Exchange of Fluid Volume Through the Capillary
Membrane 189 Starling Equilibrium for Capillary Exchange 189
Lymphatic System 190 Lymph Channels of the Body 190 Formation of
Lymph 191 Rate of Lymph Flow 192 Role of the Lymphatic System in
Controlling Interstitial Fluid Protein Concentration, Interstitial
Fluid Volume, and Interstitial Fluid Pressure 193 C H A P T E R 1 7
Local and Humoral Control of Blood Flow by the Tissues 195 Local
Control of Blood Flow in Response to Tissue Needs 195 Mechanisms of
Blood Flow Control 196 Acute Control of Local Blood Flow 196
Long-Term Blood Flow Regulation 200 Development of Collateral
CirculationA Phenomenon of Long-Term Local Blood Flow Regulation
201 Humoral Control of the Circulation 201 Vasoconstrictor Agents
201 Vasodilator Agents 202 Vascular Control by Ions and Other
Chemical Factors 202
15. Table of Contents xvii C H A P T E R 1 8 Nervous Regulation
of the Circulation, and Rapid Control of Arterial Pressure 204
Nervous Regulation of the Circulation 204 Autonomic Nervous System
204 Role of the Nervous System in Rapid Control of Arterial
Pressure 208 Increase in Arterial Pressure During Muscle Exercise
and Other Types of Stress 208 Reex Mechanisms for Maintaining
Normal Arterial Pressure 209 Central Nervous System Ischemic
ResponseControl of Arterial Pressure by the Brains Vasomotor Center
in Response to Diminished Brain Blood Flow 212 Special Features of
Nervous Control of Arterial Pressure 213 Role of the Skeletal
Nerves and Skeletal Muscles in Increasing Cardiac Output and
Arterial Pressure 213 Respiratory Waves in the Arterial Pressure
214 Arterial Pressure Vasomotor Waves Oscillation of Pressure Reex
Control Systems 214 C H A P T E R 1 9 Dominant Role of the Kidney
in Long- Term Regulation of Arterial Pressure and in Hypertension:
The Integrated System for Pressure Control 216 RenalBody Fluid
System for Arterial Pressure Control 216 Quantitation of Pressure
Diuresis as a Basis for Arterial Pressure Control 217 Chronic
Hypertension (High Blood Pressure) Is Caused by Impaired Renal
Fluid Excretion 220 The Renin-Angiotensin System: Its Role in
Pressure Control and in Hypertension 223 Components of the
Renin-Angiotensin System 223 Types of Hypertension in Which
Angiotensin Is Involved: Hypertension Caused by a Renin-Secreting
Tumor or by Infusion of Angiotensin II 226 Other Types of
Hypertension Caused by Combinations of Volume Loading and
Vasoconstriction 227 Primary (Essential) Hypertension 228 Summary
of the Integrated, Multifaceted System for Arterial Pressure
Regulation 230 C H A P T E R 2 0 Cardiac Output, Venous Return, and
Their Regulation 232 Normal Values for Cardiac Output at Rest and
During Activity 232 Control of Cardiac Output by Venous ReturnRole
of the Frank-Starling Mechanism of the Heart 232 Cardiac Output
Regulation Is the Sum of Blood Flow Regulation in All the Local
Tissues of the BodyTissue Metabolism Regulates Most Local Blood
Flow 233 The Heart Has Limits for the Cardiac Output That It Can
Achieve 234 What Is the Role of the Nervous System in Controlling
Cardiac Output? 235 Pathologically High and Pathologically Low
Cardiac Outputs 236 High Cardiac Output Caused by Reduced Total
Peripheral Resistance 236 Low Cardiac Output 237 A More
Quantitative Analysis of Cardiac Output Regulation 237 Cardiac
Output Curves Used in the Quantitative Analysis 237 Venous Return
Curves 238 Analysis of Cardiac Output and Right Atrial Pressure,
Using Simultaneous Cardiac Output and Venous Return Curves 241
Methods for Measuring Cardiac Output 243 Pulsatile Output of the
Heart as Measured by an Electromagnetic or Ultrasonic Flowmeter 243
Measurement of Cardiac Output Using the Oxygen Fick Principle 244
Indicator Dilution Method for Measuring Cardiac Output 244 C H A P
T E R 2 1 Muscle Blood Flow and Cardiac Output During Exercise; the
Coronary Circulation and Ischemic Heart Disease 246 Blood Flow in
Skeletal Muscle and Blood Flow Regulation During Exercise 246 Rate
of Blood Flow Through the Muscles 246 Control of Blood Flow Through
the Skeletal Muscles 247 Total Body Circulatory Readjustments
During Exercise 247 Coronary Circulation 249 Physiologic Anatomy of
the Coronary Blood Supply 249 Normal Coronary Blood Flow 249
Control of Coronary Blood Flow 250 Special Features of Cardiac
Muscle Metabolism 251 Ischemic Heart Disease 252 Causes of Death
After Acute Coronary Occlusion 253 Stages of Recovery from Acute
Myocardial Infarction 254 Function of the Heart After Recovery from
Myocardial Infarction 255 Pain in Coronary Heart Disease 255
Surgical Treatment of Coronary Disease 256 C H A P T E R 2 2
Cardiac Failure 258 Dynamics of the Circulation in Cardiac Failure
258
16. xviii Table of Contents Acute Effects of Moderate Cardiac
Failure 258 Chronic Stage of FailureFluid Retention Helps to
Compensate Cardiac Output 259 Summary of the Changes That Occur
After Acute Cardiac FailureCompensated Heart Failure 260 Dynamics
of Severe Cardiac Failure Decompensated Heart Failure 260
Unilateral Left Heart Failure 262 Low-Output Cardiac Failure
Cardiogenic Shock 262 Edema in Patients with Cardiac Failure 263
Cardiac Reserve 264 Quantitative Graphical Method for Analysis of
Cardiac Failure 265 C H A P T E R 2 3 Heart Valves and Heart
Sounds; Dynamics of Valvular and Congenital Heart Defects 269 Heart
Sounds 269 Normal Heart Sounds 269 Valvular Lesions 271 Abnormal
Circulatory Dynamics in Valvular Heart Disease 272 Dynamics of the
Circulation in Aortic Stenosis and Aortic Regurgitation 272
Dynamics of Mitral Stenosis and Mitral Regurgitation 273
Circulatory Dynamics During Exercise in Patients with Valvular
Lesions 273 Abnormal Circulatory Dynamics in Congenital Heart
Defects 274 Patent Ductus ArteriosusA Left-to-Right Shunt 274
Tetralogy of FallotA Right-to-Left Shunt 274 Causes of Congenital
Anomalies 276 Use of Extracorporeal Circulation During Cardiac
Surgery 276 Hypertrophy of the Heart in Valvular and Congenital
Heart Disease 276 C H A P T E R 2 4 Circulatory Shock and
Physiology of Its Treatment 278 Physiologic Causes of Shock 278
Circulatory Shock Caused by Decreased Cardiac Output 278
Circulatory Shock That Occurs Without Diminished Cardiac Output 278
What Happens to the Arterial Pressure in Circulatory Shock? 279
Tissue Deterioration Is the End Result of Circulatory Shock,
Whatever the Cause 279 Stages of Shock 279 Shock Caused by
Hypovolemia Hemorrhagic Shock 279 Relationship of Bleeding Volume
to Cardiac Output and Arterial Pressure 279 Progressive and
Nonprogressive Hemorrhagic Shock 280 Irreversible Shock 284
Hypovolemic Shock Caused by Plasma Loss 284 Hypovolemic Shock
Caused by Trauma 285 Neurogenic ShockIncreased Vascular Capacity
285 Anaphylactic Shock and Histamine Shock 285 Septic Shock 286
Physiology of Treatment in Shock 286 Replacement Therapy 286
Treatment of Shock with Sympathomimetic DrugsSometimes Useful,
Sometimes Not 287 Other Therapy 287 Circulatory Arrest 287 Effect
of Circulatory Arrest on the Brain 287 U N I T V The Body Fluids
and Kidneys C H A P T E R 2 5 The Body Fluid Compartments:
Extracellular and Intracellular Fluids; Interstitial Fluid and
Edema 291 Fluid Intake and Output Are Balanced During Steady-State
Conditions 291 Daily Intake of Water 291 Daily Loss of Body Water
291 Body Fluid Compartments 292 Intracellular Fluid Compartment 293
Extracellular Fluid Compartment 293 Blood Volume 293 Constituents
of Extracellular and Intracellular Fluids 293 Ionic Composition of
Plasma and Interstitial Fluid Is Similar 293 Important Constituents
of the Intracellular Fluid 295 Measurement of Fluid Volumes in the
Different Body Fluid Compartments The Indicator-Dilution Principle
295 Determination of Volumes of Specic Body Fluid Compartments 295
Regulation of Fluid Exchange and Osmotic Equilibrium Between
Intracellular and Extracellular Fluid 296 Basic Principles of
Osmosis and Osmotic Pressure 296 Osmotic Equilibrium Is Maintained
Between Intracellular and Extracellular Fluids 298 Volume and
Osmolality of Extracellular and Intracellular Fluids in Abnormal
States 299 Effect of Adding Saline Solution to the Extracellular
Fluid 299 Glucose and Other Solutions Administered for Nutritive
Purposes 301 Clinical Abnormalities of Fluid Volume Regulation:
Hyponatremia and Hypernatremia 301 Causes of Hyponatremia: Excess
Water or Loss of Sodium 301 Causes of Hypernatremia: Water Loss or
Excess Sodium 302 Edema: Excess Fluid in the Tissues 302
Intracellular Edema 302 Extracellular Edema 302
17. Table of Contents xix Summary of Causes of Extracellular
Edema 303 Safety Factors That Normally Prevent Edema 304 Fluids in
the Potential Spaces of the Body 305 C H A P T E R 2 6 Urine
Formation by the Kidneys: I. Glomerular Filtration, Renal Blood
Flow, and Their Control 307 Multiple Functions of the Kidneys in
Homeostasis 307 Physiologic Anatomy of the Kidneys 308 General
Organization of the Kidneys and Urinary Tract 308 Renal Blood
Supply 309 The Nephron Is the Functional Unit of the Kidney 310
Micturition 311 Physiologic Anatomy and Nervous Connections of the
Bladder 311 Transport of Urine from the Kidney Through the Ureters
and into the Bladder 312 Innervation of the Bladder 312 Filling of
the Bladder and Bladder Wall Tone; the Cystometrogram 312
Micturition Reex 313 Facilitation or Inhibition of Micturition by
the Brain 313 Abnormalities of Micturition 313 Urine Formation
Results from Glomerular Filtration, Tubular Reabsorption, and
Tubular Secretion 314 Filtration, Reabsorption, and Secretion of
Different Substances 315 Glomerular FiltrationThe First Step in
Urine Formation 316 Composition of the Glomerular Filtrate 316 GFR
Is About 20 Per Cent of the Renal Plasma Flow 316 Glomerular
Capillary Membrane 316 Determinants of the GFR 317 Increased
Glomerular Capillary Filtration Coefcient Increases GFR 318
Increased Bowmans Capsule Hydrostatic Pressure Decreases GFR 318
Increased Glomerular Capillary Colloid Osmotic Pressure Decreases
GFR 318 Increased Glomerular Capillary Hydrostatic Pressure
Increases GFR 319 Renal Blood Flow 320 Renal Blood Flow and Oxygen
Consumption 320 Determinants of Renal Blood Flow 320 Blood Flow in
the Vasa Recta of the Renal Medulla Is Very Low Compared with Flow
in the Renal Cortex 321 Physiologic Control of Glomerular
Filtration and Renal Blood Flow 321 Sympathetic Nervous System
Activation Decreases GFR 321 Hormonal and Autacoid Control of Renal
Circulation 322 Autoregulation of GFR and Renal Blood Flow 323
Importance of GFR Autoregulation in Preventing Extreme Changes in
Renal Excretion 323 Role of Tubuloglomerular Feedback in
Autoregulation of GFR 323 Myogenic Autoregulation of Renal Blood
Flow and GFR 325 Other Factors That Increase Renal Blood Flow and
GFR: High Protein Intake and Increased Blood Glucose 325 C H A P T
E R 2 7 Urine Formation by the Kidneys: II. Tubular Processing of
the Glomerular Filtrate 327 Reabsorption and Secretion by the Renal
Tubules 327 Tubular Reabsorption Is Selective and Quantitatively
Large 327 Tubular Reabsorption Includes Passive and Active
Mechanisms 328 Active Transport 328 Passive Water Reabsorption by
Osmosis Is Coupled Mainly to Sodium Reabsorption 332 Reabsorption
of Chloride, Urea, and Other Solutes by Passive Diffusion 332
Reabsorption and Secretion Along Different Parts of the Nephron 333
Proximal Tubular Reabsorption 333 Solute and Water Transport in the
Loop of Henle 334 Distal Tubule 336 Late Distal Tubule and Cortical
Collecting Tubule 336 Medullary Collecting Duct 337 Summary of
Concentrations of Different Solutes in the Different Tubular
Segments 338 Regulation of Tubular Reabsorption 339
Glomerulotubular BalanceThe Ability of the Tubules to Increase
Reabsorption Rate in Response to Increased Tubular Load 339
Peritubular Capillary and Renal Interstitial Fluid Physical Forces
339 Effect of Arterial Pressure on Urine OutputThe
Pressure-Natriuresis and Pressure-Diuresis Mechanisms 341 Hormonal
Control of Tubular Reabsorption 342 Sympathetic Nervous System
Activation Increases Sodium Reabsorption 343 Use of Clearance
Methods to Quantify Kidney Function 343 Inulin Clearance Can Be
Used to Estimate GFR 344 Creatine Clearance and Plasma Creatinine
Clearance Can Be Used to Estimate GFR 344 PAH Clearance Can Be Used
to Estimate Renal Plasma Flow 345 Filtration Fraction Is Calculated
from GFR Divided by Renal Plasma Flow 346 Calculation of Tubular
Reabsorption or Secretion from Renal Clearance 346
18. xx Table of Contents C H A P T E R 2 8 Regulation of
Extracellular Fluid Osmolarity and Sodium Concentration 348 The
Kidneys Excrete Excess Water by Forming a Dilute Urine 348
Antidiuretic Hormone Controls Urine Concentration 348 Renal
Mechanisms for Excreting a Dilute Urine 349 The Kidneys Conserve
Water by Excreting a Concentrated Urine 350 Obligatory Urine Volume
350 Requirements for Excreting a Concentrated UrineHigh ADH Levels
and Hyperosmotic Renal Medulla 350 Countercurrent Mechanism
Produces a Hyperosmotic Renal Medullary Interstitium 351 Role of
Distal Tubule and Collecting Ducts in Excreting a Concentrated
Urine 352 Urea Contributes to Hyperosmotic Renal Medullary
Interstitium and to a Concentrated Urine 353 Countercurrent
Exchange in the Vasa Recta Preserves Hyperosmolarity of the Renal
Medulla 354 Summary of Urine Concentrating Mechanism and Changes in
Osmolarity in Different Segments of the Tubules 355 Quantifying
Renal Urine Concentration and Dilution: Free Water and Osmolar
Clearances 357 Disorders of Urinary Concentrating Ability 357
Control of Extracellular Fluid Osmolarity and Sodium Concentration
358 Estimating Plasma Osmolarity from Plasma Sodium Concentration
358 Osmoreceptor-ADH Feedback System 358 ADH Synthesis in
Supraoptic and Paraventricular Nuclei of the Hypothalamus and ADH
Release from the Posterior Pituitary 359 Cardiovascular Reex
Stimulation of ADH Release by Decreased Arterial Pressure and/or
Decreased Blood Volume 360 Quantitative Importance of
Cardiovascular Reexes and Osmolarity in Stimulating ADH Secretion
360 Other Stimuli for ADH Secretion 360 Role of Thirst in
Controlling Extracellular Fluid Osmolarity and Sodium Concentration
361 Central Nervous System Centers for Thirst 361 Stimuli for
Thirst 361 Threshold for Osmolar Stimulus of Drinking 362
Integrated Responses of Osmoreceptor-ADH and Thirst Mechanisms in
Controlling Extracellular Fluid Osmolarity and Sodium Concentration
362 Role of Angiotensin II and Aldosterone in Controlling
Extracellular Fluid Osmolarity and Sodium Concentration 362
Salt-Appetite Mechanism for Controlling Extracellular Fluid Sodium
Concentration and Volume 363 C H A P T E R 2 9 Renal Regulation of
Potassium, Calcium, Phosphate, and Magnesium; Integration of Renal
Mechanisms for Control of Blood Volume and Extracellular Fluid
Volume 365 Regulation of Potassium Excretion and Potassium
Concentration in Extracellular Fluid 365 Regulation of Internal
Potassium Distribution 366 Overview of Renal Potassium Excretion
367 Potassium Secretion by Principal Cells of Late Distal and
Cortical Collecting Tubules 367 Summary of Factors That Regulate
Potassium Secretion: Plasma Potassium Concentration, Aldosterone,
Tubular Flow Rate, and Hydrogen Ion Concentration 368 Control of
Renal Calcium Excretion and Extracellular Calcium Ion Concentration
371 Control of Calcium Excretion by the Kidneys 372 Regulation of
Renal Phosphate Excretion 372 Control of Renal Magnesium Excretion
and Extracellular Magnesium Ion Concentration 373 Integration of
Renal Mechanisms for Control of Extracellular Fluid 373 Sodium
Excretion Is Precisely Matched to Intake Under Steady-State
Conditions 373 Sodium Excretion Is Controlled by Altering
Glomerular Filtration or Tubular Sodium Reabsorption Rates 374
Importance of Pressure Natriuresis and Pressure Diuresis in
Maintaining Body Sodium and Fluid Balance 374 Pressure Natriuresis
and Diuresis Are Key Components of a Renal-Body Fluid Feedback for
Regulating Body Fluid Volumes and Arterial Pressure 375 Precision
of Blood Volume and Extracellular Fluid Volume Regulation 376
Distribution of Extracellular Fluid Between the Interstitial Spaces
and Vascular System 376 Nervous and Hormonal Factors Increase the
Effectiveness of Renal-Body Fluid Feedback Control 377 Sympathetic
Nervous System Control of Renal Excretion: Arterial Baroreceptor
and Low-Pressure Stretch Receptor Reexes 377 Role of Angiotensin II
In Controlling Renal Excretion 377 Role of Aldosterone in
Controlling Renal Excretion 378 Role of ADH in Controlling Renal
Water Excretion 379 Role of Atrial Natriuretic Peptide in
Controlling Renal Excretion 378 Integrated Responses to Changes in
Sodium Intake 380 Conditions That Cause Large Increases in Blood
Volume and Extracellular Fluid Volume 380
19. Table of Contents xxi Increased Blood Volume and
Extracellular Fluid Volume Caused by Heart Diseases 380 Increased
Blood Volume Caused by Increased Capacity of Circulation 380
Conditions That Cause Large Increases in Extracellular Fluid Volume
but with Normal Blood Volume 381 Nephrotic SyndromeLoss of Plasma
Proteins in Urine and Sodium Retention by the Kidneys 381 Liver
CirrhosisDecreased Synthesis of Plasma Proteins by the Liver and
Sodium Retention by the Kidneys 381 C H A P T E R 3 0 Regulation of
Acid-Base Balance 383 Hydrogen Ion Concentration Is Precisely
Regulated 383 Acids and BasesTheir Denitions and Meanings 383
Defenses Against Changes in Hydrogen Ion Concentration: Buffers,
Lungs, and Kidneys 384 Buffering of Hydrogen Ions in the Body
Fluids 385 Bicarbonate Buffer System 385 Quantitative Dynamics of
the Bicarbonate Buffer System 385 Phosphate Buffer System 387
Proteins: Important Intracellular Buffers 387 Respiratory
Regulation of Acid-Base Balance 388 Pulmonary Expiration of CO2
Balances Metabolic Formation of CO2 388 Increasing Alveolar
Ventilation Decreases Extracellular Fluid Hydrogen Ion
Concentration and Raises pH 388 Increased Hydrogen Ion
Concentration Stimulates Alveolar Ventilation 389 Renal Control of
Acid-Base Balance 390 Secretion of Hydrogen Ions and Reabsorption
of Bicarbonate Ions by the Renal Tubules 390 Hydrogen Ions Are
Secreted by Secondary Active Transport in the Early Tubular
Segments 391 Filtered Bicarbonate Ions Are Reabsorbed by
Interaction with Hydrogen Ions in the Tubules 391 Primary Active
Secretion of Hydrogen Ions in the Intercalated Cells of Late Distal
and Collecting Tubules 392 Combination of Excess Hydrogen Ions with
Phosphate and Ammonia Buffers in the TubuleA Mechanism for
Generating New Bicarbonate Ions 392 Phosphate Buffer System Carries
Excess Hydrogen Ions into the Urine and Generates New Bicarbonate
393 Excretion of Excess Hydrogen Ions and Generation of New
Bicarbonate by the Ammonia Buffer System 393 Quantifying Renal
Acid-Base Excretion 394 Regulation of Renal Tubular Hydrogen Ion
Secretion 395 Renal Correction of AcidosisIncreased Excretion of
Hydrogen Ions and Addition of Bicarbonate Ions to the Extracellular
Fluid 396 Acidosis Decreases the Ratio of HCO3 - /H+ in Renal
Tubular Fluid 396 Renal Correction of AlkalosisDecreased Tubular
Secretion of Hydrogen Ions and Increased Excretion of Bicarbonate
Ions 396 Alkalosis Increases the Ratio of HCO3 - /H+ in Renal
Tubular Fluid 396 Clinical Causes of Acid-Base Disorders 397
Respiratory Acidosis Is Caused by Decreased Ventilation and
Increased PCO2 397 Respiratory Alkalosis Results from Increased
Ventilation and Decreased PCO2 397 Metabolic Acidosis Results from
Decreased Extracellular Fluid Bicarbonate Concentration 397
Treatment of Acidosis or Alkalosis 398 Clinical Measurements and
Analysis of Acid-Base Disorders 398 Complex Acid-Base Disorders and
Use of the Acid-Base Nomogram for Diagnosis 399 Use of Anion Gap to
Diagnose Acid-Base Disorders 400 C H A P T E R 3 1 Kidney Diseases
and Diuretics 402 Diuretics and Their Mechanisms of Action 402
Osmotic Diuretics Decrease Water Reabsorption by Increasing Osmotic
Pressure of Tubular Fluid 402 Loop Diuretics Decrease Active
Sodium-Chloride-Potassium Reabsorption in the Thick Ascending Loop
of Henle 403 Thiazide Diuretics Inhibit Sodium-Chloride
Reabsorption in the Early Distal Tubule 404 Carbonic Anhydrase
Inhibitors Block Sodium-Bicarbonate Reabsorption in the Proximal
Tubules 404 Competitive Inhibitors of Aldosterone Decrease Sodium
Reabsorption from and Potassium Secretion into the Cortical
Collecting Tubule 404 Diuretics That Block Sodium Channels in the
Collecting Tubules Decrease Sodium Reabsorption 404 Kidney Diseases
404 Acute Renal Failure 404 Prerenal Acute Renal Failure Caused by
Decreased Blood Flow to the Kidney 405 Intrarenal Acute Renal
Failure Caused by Abnormalities within the Kidney 405 Postrenal
Acute Renal Failure Caused by Abnormalities of the Lower Urinary
Tract 406 Physiologic Effects of Acute Renal Failure 406 Chronic
Renal Failure: An Irreversible Decrease in the Number of Functional
Nephrons 406 Vicious Circle of Chronic Renal Failure Leading to
End-Stage Renal Disease 407 Injury to the Renal Vasculature as a
Cause of Chronic Renal Failure 408
20. xxii Table of Contents Injury to the Glomeruli as a Cause
of Chronic Renal Failure Glomerulonephritis 408 Injury to the Renal
Interstitium as a Cause of Chronic Renal Failure Pyelonephritis 409
Nephrotic SyndromeExcretion of Protein in the Urine Because of
Increased Glomerular Permeability 409 Nephron Function in Chronic
Renal Failure 409 Effects of Renal Failure on the Body FluidsUremia
411 Hypertension and Kidney Disease 412 Specic Tubular Disorders
413 Treatment of Renal Failure by Dialysis with an Articial Kidney
414 U N I T V I Blood Cells, Immunity, and Blood Clotting C H A P T
E R 3 2 Red Blood Cells, Anemia, and Polycythemia 419 Red Blood
Cells (Erythrocytes) 419 Production of Red Blood Cells 420
Formation of Hemoglobin 424 Iron Metabolism 425 Life Span and
Destruction of Red Blood Cells 426 Anemias 426 Effects of Anemia on
Function of the Circulatory System 427 Polycythemia 427 Effect of
Polycythemia on Function of the Circulatory System 428 C H A P T E
R 3 3 Resistance of the Body to Infection: I. Leukocytes,
Granulocytes, the Monocyte-Macrophage System, and Inammation 429
Leukocytes (White Blood Cells) 429 General Characteristics of
Leukocytes 429 Genesis of the White Blood Cells 430 Life Span of
the White Blood Cells 431 Neutrophils and Macrophages Defend
Against Infections 431 Phagocytosis 431 Monocyte-Macrophage Cell
System (Reticuloendothelial System) 432 Inammation: Role of
Neutrophils and Macrophages 434 Inammation 434 Macrophage and
Neutrophil Responses During Inammation 434 Eosinophils 436
Basophils 436 Leukopenia 436 The Leukemias 437 Effects of Leukemia
on the Body 437 C H A P T E R 3 4 Resistance of the Body to
Infection: II. Immunity and Allergy 439 Innate Immunity 439
Acquired (Adaptive) Immunity 439 Basic Types of Acquired Immunity
440 Both Types of Acquired Immunity Are Initiated by Antigens 440
Lymphocytes Are Responsible for Acquired Immunity 440 Preprocessing
of the T and B Lymphocytes 440 T Lymphocytes and B-Lymphocyte
Antibodies React Highly Specically Against Specic AntigensRole of
Lymphocyte Clones 442 Origin of the Many Clones of Lymphocytes 442
Specic Attributes of the B-Lymphocyte SystemHumoral Immunity and
the Antibodies 443 Special Attributes of the T-Lymphocyte
SystemActivated T Cells and Cell- Mediated Immunity 446 Several
Types of T Cells and Their Different Functions 446 Tolerance of the
Acquired Immunity System to Ones Own TissuesRole of Preprocessing
in the Thymus and Bone Marrow 448 Immunization by Injection of
Antigens 448 Passive Immunity 449 Allergy and Hypersensitivity 449
Allergy Caused by Activated T Cells: Delayed-Reaction Allergy 449
Allergies in the Allergic Person, Who Has Excess IgE Antibodies 449
C H A P T E R 3 5 Blood Types; Transfusion; Tissue and Organ
Transplantation 451 Antigenicity Causes Immune Reactions of Blood
451 O-A-B Blood Types 451 A and B AntigensAgglutinogens 451
Agglutinins 452 Agglutination Process In Transfusion Reactions 452
Blood Typing 453 Rh Blood Types 453 Rh Immune Response 453
Transfusion Reactions Resulting from Mismatched Blood Types 454
Transplantation of Tissues and Organs 455 Attempts to Overcome
Immune Reactions in Transplanted Tissue 455 C H A P T E R 3 6
Hemostasis and Blood Coagulation 457 Events in Hemostasis 457
Vascular Constriction 457 Formation of the Platelet Plug 457 Blood
Coagulation in the Ruptured Vessel 458 Fibrous Organization or
Dissolution of the Blood Clot 458
21. Table of Contents xxiii Mechanism of Blood Coagulation 459
Conversion of Prothrombin to Thrombin 459 Conversion of Fibrinogen
to Fibrin Formation of the Clot 460 Vicious Circle of Clot
Formation 460 Initiation of Coagulation: Formation of Prothrombin
Activator 461 Prevention of Blood Clotting in the Normal Vascular
SystemIntravascular Anticoagulants 463 Lysis of Blood ClotsPlasmin
464 Conditions That Cause Excessive Bleeding in Human Beings 464
Decreased Prothrombin, Factor VII, Factor IX,and Factor X Caused by
Vitamin K Deciency 464 Hemophilia 465 Thrombocytopenia 465
Thromboembolic Conditions in the Human Being 465 Femoral Venous
Thrombosis and Massive Pulmonary Embolism 466 Disseminated
Intravascular Coagulation 466 Anticoagulants for Clinical Use 466
Heparin as an Intravenous Anticoagulant 466 Coumarins as
Anticoagulants 466 Prevention of Blood Coagulation Outside the Body
466 Blood Coagulation Tests 467 Bleeding Time 467 Clotting Time 467
Prothrombin Time 467 U N I T V I I Respiration C H A P T E R 3 7
Pulmonary Ventilation 471 Mechanics of Pulmonary Ventilation 471
Muscles That Cause Lung Expansion and Contraction 471 Movement of
Air In and Out of the Lungs and the Pressures That Cause the
Movement 472 Effect of the Thoracic Cage on Lung Expansibility 474
Pulmonary Volumes and Capacities 475 Recording Changes in Pulmonary
Volume Spirometry 475 Abbreviations and Symbols Used in Pulmonary
Function Tests 476 Determination of Functional Residual Capacity,
Residual Volume, and Total Lung CapacityHelium Dilution Method 476
Minute Respiratory Volume Equals Respiratory Rate Times Tidal
Volume 477 Alveolar Ventilation 477 Dead Space and Its Effect on
Alveolar Ventilation 477 Rate of Alveolar Ventilation 478 Functions
of the Respiratory Passageways 478 Trachea, Bronchi, and
Bronchioles 478 Normal Respiratory Functions of the Nose 480 C H A
P T E R 3 8 Pulmonary Circulation, Pulmonary Edema, Pleural Fluid
483 Physiologic Anatomy of the Pulmonary Circulatory System 483
Pressures in the Pulmonary System 483 Blood Volume of the Lungs 484
Blood Flow Through the Lungs and Its Distribution 485 Effect of
Hydrostatic Pressure Gradients in the Lungs on Regional Pulmonary
Blood Flow 485 Zones 1, 2, and 3 of Pulmonary Blood Flow 485 Effect
of Increased Cardiac Output on Pulmonary Blood Flow and Pulmonary
Arterial Pressure During Heavy Exercise 486 Function of the
Pulmonary Circulation When the Left Atrial Pressure Rises as a
Result of Left-Sided Heart Failure 487 Pulmonary Capillary Dynamics
487 Capillary Exchange of Fluid in the Lungs, and Pulmonary
Interstitial Fluid Dynamics 487 Pulmonary Edema 488 Fluid in the
Pleural Cavity 489 C H A P T E R 3 9 Physical Principles of Gas
Exchange; Diffusion of Oxygen and Carbon Dioxide Through the
Respiratory Membrane 491 Physics of Gas Diffusion and Gas Partial
Pressures 491 Molecular Basis of Gas Diffusion 491 Gas Pressures in
a Mixture of Gases Partial Pressures of Individual Gases 491
Pressures of Gases Dissolved in Water and Tissues 492 Vapor
Pressure of Water 492 Diffusion of Gases Through Fluids Pressure
Difference Causes Net Diffusion 493 Diffusion of Gases Through
Tissues 493 Composition of Alveolar AirIts Relation to Atmospheric
Air 493 Rate at Which Alveolar Air Is Renewed by Atmospheric Air
494 Oxygen Concentration and Partial Pressure in the Alveoli 494
CO2 Concentration and Partial Pressure in the Alveoli 495 Expired
Air 495 Diffusion of Gases Through the Respiratory Membrane 496
Factors That Affect the Rate of Gas Diffusion Through the
Respiratory Membrane 498 Diffusing Capacity of the Respiratory
Membrane 498 Effect of the Ventilation-Perfusion Ratio on Alveolar
Gas Concentration 499 PO2-PCO2, V . A/Q . Diagram 500 Concept of
the Physiological Shunt (When V . A/Q . Is Greater Than Normal) 500
Abnormalities of Ventilation-Perfusion Ratio 501
22. xxiv Table of Contents C H A P T E R 4 0 Transport of
Oxygen and Carbon Dioxide in Blood and Tissue Fluids 502 Transport
of Oxygen from the Lungs to the Body Tissues 502 Diffusion of
Oxygen from the Alveoli to the Pulmonary Capillary Blood 502
Transport of Oxygen in the Arterial Blood 503 Diffusion of Oxygen
from the Peripheral Capillaries into the Tissue Fluid 503 Diffusion
of Oxygen from the Peripheral Capillaries to the Tissue Cells 504
Diffusion of Carbon Dioxide from the Peripheral Tissue Cells into
the Capillaries and from the Pulmonary Capillaries into the Alveoli
504 Role of Hemoglobin in Oxygen Transport 505 Reversible
Combination of Oxygen with Hemoglobin 505 Effect of Hemoglobin to
Buffer the Tissue PO2 507 Factors That Shift the Oxygen-Hemoglobin
Dissociation CurveTheir Importance for Oxygen Transport 507
Metabolic Use of Oxygen by the Cells 508 Transport of Oxygen in the
Dissolved State 509 Combination of Hemoglobin with Carbon
MonoxideDisplacement of Oxygen 509 Transport of Carbon Dioxide in
the Blood 510 Chemical Forms in Which Carbon Dioxide Is Transported
510 Carbon Dioxide Dissociation Curve 511 When Oxygen Binds with
Hemoglobin, Carbon Dioxide Is Released (the Haldane Effect) to
Increase CO2 Transport 511 Change in Blood Acidity During Carbon
Dioxide Transport 512 Respiratory Exchange Ratio 512 C H A P T E R
4 1 Regulation of Respiration 514 Respiratory Center 514 Dorsal
Respiratory Group of NeuronsIts Control of Inspiration and of
Respiratory Rhythm 514 A Pneumotaxic Center Limits the Duration of
Inspiration and Increases the Respiratory Rate 514 Ventral
Respiratory Group of Neurons Functions in Both Inspiration and
Expiration 515 Lung Ination Signals Limit Inspiration The
Hering-Breuer Ination Reex 515 Control of Overall Respiratory
Center Activity 516 Chemical Control of Respiration 516 Direct
Chemical Control of Respiratory Center Activity by Carbon Dioxide
and Hydrogen Ions 516 Peripheral Chemoreceptor System for Control
of Respiratory ActivityRole of Oxygen in Respiratory Control 518
Effect of Low Arterial PO2 to Stimulate Alveolar Ventilation When
Arterial Carbon Dioxide and Hydrogen Ion Concentrations Remain
Normal 519 Chronic Breathing of Low Oxygen Stimulates Respiration
Even MoreThe Phenomenon of Acclimatization 519 Composite Effects of
PCO2, pH, and PO2 on Alveolar Ventilation 519 Regulation of
Respiration During Exercise 520 Other Factors That Affect
Respiration 521 Sleep Apnea 522 C H A P T E R 4 2 Respiratory
Insufciency Pathophysiology, Diagnosis, Oxygen Therapy 524 Useful
Methods for Studying Respiratory Abnormalities 524 Study of Blood
Gases and Blood pH 524 Measurement of Maximum Expiratory Flow 525
Forced Expiratory Vital Capacity and Forced Expiratory Volume 526
Physiologic Peculiarities of Specic Pulmonary Abnormalities 526
Chronic Pulmonary Emphysema 526 Pneumonia 527 Atelectasis 528
Asthma 529 Tuberculosis 530 Hypoxia and Oxygen Therapy 530 Oxygen
Therapy in Different Types of Hypoxia 530 Cyanosis 531 Hypercapnia
531 Dyspnea 532 Articial Respiration 532 U N I T V I I I Aviation,
Space, and Deep-Sea Diving Physiology C H A P T E R 4 3 Aviation,
High-Altitude, and Space Physiology 537 Effects of Low Oxygen
Pressure on the Body 537 Alveolar PO2 at Different Elevations 537
Effect of Breathing Pure Oxygen on Alveolar PO2 at Different
Altitudes 538 Acute Effects of Hypoxia 538 Acclimatization to Low
PO2 539 Natural Acclimatization of Native Human Beings Living at
High Altitudes 540 Reduced Work Capacity at High Altitudes and
Positive Effect of Acclimatization 540 Acute Mountain Sickness and
High-Altitude Pulmonary Edema 540 Chronic Mountain Sickness 541
Effects of Acceleratory Forces on the Body in Aviation and Space
Physiology 541 Centrifugal Acceleratory Forces 541 Effects of
Linear Acceleratory Forces on the Body 542
23. Table of Contents xxv Articial Climate in the Sealed
Spacecraft 543 Weightlessness in Space 543 C H A P T E R 4 4
Physiology of Deep-Sea Diving and Other Hyperbaric Conditions 545
Effect of High Partial Pressures of Individual Gases on the Body
545 Nitrogen Narcosis at High Nitrogen Pressures 545 Oxygen
Toxicity at High Pressures 546 Carbon Dioxide Toxicity at Great
Depths in the Sea 547 Decompression of the Diver After Excess
Exposure to High Pressure 547 Scuba (Self-Contained Underwater
Breathing Apparatus) Diving 549 Special Physiologic Problems in
Submarines 550 Hyperbaric Oxygen Therapy 550 U N I T I X The
Nervous System: A. General Principles and Sensory Physiology C H A
P T E R 4 5 Organization of the Nervous System, Basic Functions of
Synapses, Transmitter Substances 555 General Design of the Nervous
System 555 Central Nervous System Neuron: The Basic Functional Unit
555 Sensory Part of the Nervous System Sensory Receptors 555 Motor
Part of the Nervous System Effectors 556 Processing of
InformationIntegrative Function of the Nervous System 556 Storage
of InformationMemory 557 Major Levels of Central Nervous System
Function 557 Spinal Cord Level 557 Lower Brain or Subcortical Level
558 Higher Brain or Cortical Level 558 Comparison of the Nervous
System with a Computer 558 Central Nervous System Synapses 559
Types of SynapsesChemical and Electrical 559 Physiologic Anatomy of
the Synapse 559 Chemical Substances That Function as Synaptic
Transmitters 562 Electrical Events During Neuronal Excitation 564
Electrical Events During Neuronal Inhibition 566 Special Functions
of Dendrites for Exciting Neurons 568 Relation of State of
Excitation of the Neuron to Rate of Firing 569 Some Special
Characteristics of Synaptic Transmission 570 C H A P T E R 4 6
Sensory Receptors, Neuronal Circuits for Processing Information 572
Types of Sensory Receptors and the Sensory Stimuli They Detect 572
Differential Sensitivity of Receptors 572 Transduction of Sensory
Stimuli into Nerve Impulses 573 Local Electrical Currents at Nerve
Endings Receptor Potentials 573 Adaptation of Receptors 575 Nerve
Fibers That Transmit Different Types of Signals, and Their
Physiologic Classication 576 Transmission of Signals of Different
Intensity in Nerve TractsSpatial and Temporal Summation 577
Transmission and Processing of Signals in Neuronal Pools 578
Relaying of Signals Through Neuronal Pools 579 Prolongation of a
Signal by a Neuronal PoolAfterdischarge 581 Instability and
Stability of Neuronal Circuits 583 Inhibitory Circuits as a
Mechanism for Stabilizing Nervous System Function 583 Synaptic
Fatigue as a Means for Stabilizing the Nervous System 583 C H A P T
E R 4 7 Somatic Sensations: I. General Organization, the Tactile
and Position Senses 585 CLASSIFICATION OF SOMATIC SENSES 585
Detection and Transmission of Tactile Sensations 585 Detection of
Vibration 587 TICKLE AND ITCH 587 Sensory Pathways for Transmitting
Somatic Signals into the Central Nervous System 587 Dorsal
ColumnMedial Lemniscal System 588 Anterolateral System 588
Transmission in the Dorsal Column Medial Lemniscal System 588
Anatomy of the Dorsal ColumnMedial Lemniscal System 588
Somatosensory Cortex 589 Somatosensory Association Areas 592
Overall Characteristics of Signal Transmission and Analysis in the
Dorsal ColumnMedial Lemniscal System 592 Position Senses 594
Interpretation of Sensory Stimulus Intensity 593 Judgment of
Stimulus Intensity 594 Position Senses 594 Transmission of Less
Critical Sensory Signals in the Anterolateral Pathway 595 Anatomy
of the Anterolateral Pathway 595 Some Special Aspects of
Somatosensory Function 596 Function of the Thalamus in Somatic
Sensation 596
24. xxvi Table of Contents Cortical Control of Sensory
Sensitivity Corticofugal Signals 597 Segmental Fields of
SensationThe Dermatomes 597 C H A P T E R 4 8 Somatic Sensations:
II. Pain, Headache, and Thermal Sensations 598 Types of Pain and
Their Qualities Fast Pain and Slow Pain 598 Pain Receptors and
Their Stimulation 598 Rate of Tissue Damage as a Stimulus for Pain
599 Dual Pathways for Transmission of Pain Signals into the Central
Nervous System 600 Dual Pain Pathways in the Cord and Brain StemThe
Neospinothalamic Tract and the Paleospinothalamic Tract 600 Pain
Suppression (Analgesia) System in the Brain and Spinal Cord 602
Brains Opiate SystemEndorphins and Enkephalins 602 Inhibition of
Pain Transmission by Simultaneous Tactile Sensory Signals 603
Treatment of Pain by Electrical Stimulation 603 Referred Pain 603
Visceral Pain 603 Causes of True Visceral Pain 604 Parietal Pain
Caused by Visceral Disease 604 Localization of Visceral
PainVisceral and the Parietal Pain Transmission Pathways 604 Some
Clinical Abnormalities of Pain and Other Somatic Sensations 605
Hyperalgesia 605 Herpes Zoster (Shingles) 605 Tic Douloureux 605
Brown-Squard Syndrome 606 Headache 606 Headache of Intracranial
Origin 606 Thermal Sensations 607 Thermal Receptors and Their
Excitation 607 Transmission of Thermal Signals in the Nervous
System 609 U N I T X The Nervous System: B. The Special Senses C H
A P T E R 4 9 The Eye: I. Optics of Vision 613 Physical Principles
of Optics 613 Refraction of Light 613 Application of Refractive
Principles to Lenses 613 Focal Length of a Lens 615 Formation of an
Image by a Convex Lens 616 Measurement of the Refractive Power of a
LensDiopter 616 Optics of the Eye 617 The Eye as a Camera 617
Mechanism of Accommodation 617 Pupillary Diameter 618 Errors of
Refraction 619 Visual Acuity 621 Determination of Distance of an
Object from the EyeDepth Perception 621 Ophthalmoscope 622 Fluid
System of the EyeIntraocular Fluid 623 Formation of Aqueous Humor
by the Ciliary Body 623 Outow of Aqueous Humor from the Eye 623
Intraocular Pressure 624 C H A P T E R 5 0 The Eye: II. Receptor
and Neural Function of the Retina 626 Anatomy and Function of the
Structural Elements of the Retina 626 Photochemistry of Vision 628
Rhodopsin-Retinal Visual Cycle, and Excitation of the Rods 629
Automatic Regulation of Retinal Sensitivity Light and Dark
Adaptation 631 Color Vision 632 Tricolor Mechanism of Color
Detection 632 Color Blindness 633 Neural Function of the Retina 633
Neural Circuitry of the Retina 633 Ganglion Cells and Optic Nerve
Fibers 636 Excitation of the Ganglion Cells 637 C H A P T E R 5 1
The Eye: III. Central Neurophysiology of Vision 640 Visual Pathways
640 Function of the Dorsal Lateral Geniculate Nucleus of the
Thalamus 640 Organization and Function of the Visual Cortex 641
Layered Structure of the Primary Visual Cortex 642 Two Major
Pathways for Analysis of Visual Information(1) The Fast Position
and Motion Pathway; (2) The Accurate Color Pathway 643 Neuronal
Patterns of Stimulation During Analysis of the Visual Image 643
Detection of Color 644 Effect of Removing the Primary Visual Cortex
644 Fields of Vision; Perimetry 644 Eye Movements and Their Control
645 Fixation Movements of the Eyes 645 Fusion of the Visual Images
from the Two Eyes 647 Autonomic Control of Accommodation and
Pupillary Aperture 648 Control of Accommodation (Focusing the Eyes)
649 Control of Pupillary Diameter 649 C H A P T E R 5 2 The Sense
of Hearing 651 Tympanic Membrane and the Ossicular System 651
25. Table of Contents xxvii Conduction of Sound from the
Tympanic Membrane to the Cochlea 651 Transmission of Sound Through
Bone 652 Cochlea 652 Functional Anatomy of the Cochlea 652
Transmission of Sound Waves in the CochleaTraveling Wave 654
Function of the Organ of Corti 655 Determination of Sound
FrequencyThe Place Principle 656 Determination of Loudness 656
Central Auditory Mechanisms 657 Auditory Nervous Pathways 657
Function of the Cerebral Cortex in Hearing 658 Determination of the
Direction from Which Sound Comes 660 Centrifugal Signals from the
Central Nervous System to Lower Auditory Centers 660 Hearing
Abnormalities 660 Types of Deafness 660 C H A P T E R 5 3 The
Chemical SensesTaste and Smell 663 Sense of Taste 663 Primary
Sensations of Taste 663 Taste Bud and Its Function 664 Transmission
of Taste Signals into the Central Nervous System 665 Taste
Preference and Control of the Diet 666 Sense of Smell 667 Olfactory
Membrane 667 Stimulation of the Olfactory Cells 667 Transmission of
Smell Signals into the Central Nervous System 668 U N I T X I The
Nervous System: C. Motor and Integrative Neurophysiology C H A P T
E R 5 4 Motor Functions of the Spinal Cord; the Cord Reexes 673
Organization of the Spinal Cord for Motor Functions 673 Muscle
Sensory ReceptorsMuscle Spindles and Golgi Tendon Organs And Their
Roles in Muscle Control 675 Receptor Function of the Muscle Spindle
675 Muscle Stretch Reex 676 Role of the Muscle Spindle in Voluntary
Motor Activity 678 Clinical Applications of the Stretch Reex 678
Golgi Tendon Reex 679 Function of the Muscle Spindles and Golgi
Tendon Organs in Conjunction with Motor Control from Higher Levels
of the Brain 680 Flexor Reex and the Withdrawal Reexes 680 Crossed
Extensor Reex 681 Reciprocal Inhibition and Reciprocal Innervation
681 Reexes of Posture and Locomotion 682 Postural and Locomotive
Reexes of the Cord 682 Scratch Reex 683 Spinal Cord Reexes That
Cause Muscle Spasm 683 Autonomic Reexes in the Spinal Cord 683
Spinal Cord Transection and Spinal Shock 684 C H A P T E R 5 5
Cortical and Brain Stem Control of Motor Function 685 MOTOR CORTEX
AND CORTICOSPINAL TRACT 685 Primary Motor Cortex 685 Premotor Area
686 Supplementary Motor Area 686 Some Specialized Areas of Motor
Control Found in the Human Motor Cortex 686 Transmission of Signals
from the Motor Cortex to the Muscles 687 Incoming Fiber Pathways to
the Motor Cortex 688 Red Nucleus Serves as an Alternative Pathway
for Transmitting Cortical Signals to the Spinal Cord 688
Extrapyramidal System 689 Excitation of the Spinal Cord Motor
Control Areas by the Primary Motor Cortex and Red Nucleus 689 Role
of the Brain Stem in Controlling Motor Function 691 Support of the
Body Against Gravity Roles of the Reticular and Vestibular Nuclei
691 Vestibular Sensations and Maintenance of Equilibrium 692
Vestibular Apparatus 692 Function of the Utricle and Saccule in the
Maintenance of Static Equilibrium 694 Detection of Head Rotation by
the Semicircular Ducts 695 Vestibular Mechanisms for Stabilizing
the Eyes 696 Other Factors Concerned with Equilibrium 696 Functions
of Brain Stem Nuclei in Controlling Subconscious, Stereotyped
Movements 697 C H A P T E R 5 6 Contributions of the Cerebellum and
Basal Ganglia to Overall Motor Control 698 Cerebellum and Its Motor
Functions 698 Anatomical Functional Areas of the Cerebellum 699
Neuronal Circuit of the Cerebellum 700 Function of the Cerebellum
in Overall Motor Control 703 Clinical Abnormalities of the
Cerebellum 706
26. xxviii Table of Contents Basal GangliaTheir Motor Functions
707 Function of the Basal Ganglia in Executing Patterns of Motor
ActivityThe Putamen Circuit 708 Role of the Basal Ganglia for
Cognitive Control of Sequences of Motor Patterns The Caudate
Circuit 709 Function of the Basal Ganglia to Change the Timing and
to Scale the Intensity of Movements 709 Functions of Specic
Neurotransmitter Substances in the Basal Ganglial System 710
Integration of the Many Parts of the Total Motor Control System 712
Spinal Level 712 Hindbrain Level 712 Motor Cortex Level 712 What
Drives Us to Action? 713 C H A P T E R 5 7 Cerebral Cortex,
Intellectual Functions of the Brain, Learning and Memory 714
Physiologic Anatomy of the Cerebral Cortex 714 Functions of Specic
Cortical Areas 715 Association Areas 716 Comprehensive
Interpretative Function of the Posterior Superior Temporal Lobe
Wernickes Area (a General Interpretative Area) 718 Functions of the
Parieto-occipitotemporal Cortex in the Nondominant Hemisphere 719
Higher Intellectual Functions of the Prefrontal Association Areas
719 Function of the Brain in CommunicationLanguage Input and
Language Output 720 Function of the Corpus Callosum and Anterior
Commissure to Transfer Thoughts, Memories, Training, and Other
Information Between the Two Cerebral Hemispheres 722 Thoughts,
Consciousness, and Memory 723 MemoryRoles of Synaptic Facilitation
and Synaptic Inhibition 723 Short-Term Memory 724 Intermediate
Long-Term Memory 724 Long-Term Memory 725 Consolidation of Memory
725 C H A P T E R 5 8 Behavioral and Motivational Mechanisms of the
BrainThe Limbic System and the Hypothalamus 728 Activating-Driving
Systems of the Brain 728 Control of Cerebral Activity by Continuous
Excitatory Signals from the Brain Stem 728 Neurohormonal Control of
Brain Activity 730 Limbic System 731 Functional Anatomy of the
Limbic System; Key Position of the Hypothalamus 731 Hypothalamus, a
Major Control Headquarters for the Limbic System 732 Vegetative and
Endocrine Control Functions of the Hypothalamus 733 Behavioral
Functions of the Hypothalamus and Associated Limbic Structures 734
Reward and Punishment Function of the Limbic System 735 Importance
of Reward or Punishment in Behavior 736 Specic Functions of Other
Parts of the Limbic System 736 Functions of the Hippocampus 736
Functions of the Amygdala 737 Function of the Limbic Cortex 738 C H
A P T E R 5 9 States of Brain ActivitySleep, Brain Waves, Epilepsy,
Psychoses 739 Sleep 739 Slow-Wave Sleep 739 REM Sleep (Paradoxical
Sleep, Desynchronized Sleep) 740 Basic Theories of Sleep 740
Physiologic Effects of Sleep 741 Brain Waves 741 Origin of Brain
Waves 742 Effect of Varying Levels of Cerebral Activity on the
Frequency of the EEG 743 Changes in the EEG at Different Stages of
Wakefulness and Sleep 743 Epilepsy 743 Grand Mal Epilepsy 743 Petit
Mal Epilepsy 744 Focal Epilepsy 744 Psychotic Behavior and Dementia
Roles of Specic Neurotransmitter Systems 745 Depression and
Manic-Depressive PsychosesDecreased Activity of the Norepinephrine
and Serotonin Neurotransmitter Systems 745 SchizophreniaPossible
Exaggerated Function of Part of the Dopamine System 745 Alzheimers
DiseaseAmyloid Plaques and Depressed Memory 746 C H A P T E R 6 0
The Autonomic Nervous System and the Adrenal Medulla 748 General
Organization of the Autonomic Nervous System 748 Physiologic
Anatomy of the Sympathetic Nervous System 748 Preganglionic and
Postganglionic Sympathetic Neurons 748 Physiologic Anatomy of the
Parasympathetic Nervous System 750 Basic Characteristics of
Sympathetic and Parasympathetic Function 750 Cholinergic and
Adrenergic Fibers Secretion of Acetylcholine or Norepinephrine 750
Receptors on the Effector Organs 752
27. Table of Contents xxix Excitatory and Inhibitory Actions of
Sympathetic and Parasympathetic Stimulation 753 Effects of
Sympathetic and Parasympathetic Stimulation on Specic Organs 753
Function of the Adrenal Medullae 755 Relation of Stimulus Rate to
Degree of Sympathetic and Parasympathetic Effect 756 Sympathetic
and Parasympathetic Tone 756 Denervation Supersensitivity of
Sympathetic and Parasympathetic Organs after Denervation 756
Autonomic Reexes 757 Stimulation of Discrete Organs in Some
Instances and Mass Stimulation in Other Instances by the
Sympathetic and Parasympathetic Systems 757 Alarm or Stress
Response of the Sympathetic Nervous System 758 Medullary, Pontine,
and Mesencephalic Control of the Autonomic Nervous System 758
Pharmacology of the Autonomic Nervous System 759 Drugs That Act on
Adrenergic Effector OrgansSympathomimetic Drugs 759 Drugs That Act
on Cholinergic Effector Organs 759 Drugs That Stimulate or Block
Sympathetic and Parasympathetic Postganglionic Neurons 759 C H A P
T E R 6 1 Cerebral Blood Flow, Cerebrospinal Fluid, and Brain
Metabolism 761 Cerebral Blood Flow 761 Normal Rate of Cerebral
Blood Flow 761 Regulation of Cerebral Blood Flow 761 Cerebral
Microcirculation 763 Cerebral Stroke Occurs When Cerebral Blood
Vessels are Blocked 763 Cerebrospinal Fluid System 763 Cushioning
Function of the Cerebrospinal Fluid 763 Formation, Flow, and
Absorption of Cerebrospinal Fluid 764 Cerebrospinal Fluid Pressure
765 Obstruction to Flow of Cerebrospinal Fluid Can Cause
Hydrocephalus 766 BloodCerebrospinal Fluid and Blood-Brain Barriers
766 Brain Edema 766 Brain Metabolism 767 U N I T X I I
Gastrointestinal Physiology C H A P T E R 6 2 General Principles of
Gastrointestinal FunctionMotility, Nervous Control, and Blood
Circulation 771 General Principles of Gastrointestinal Motility 771
Physiological Anatomy of the Gastrointestinal Wall 771 Neural
Control of Gastrointestinal FunctionEnteric Nervous System 773
Differences Between the Myenteric and Submucosal Plexuses 774 Types
of Neurotransmitters Secreted by Enteric Neurons 775 Hormonal
Control of Gastrointestinal Motility 776 Functional Types of
Movements in the Gastrointestinal Tract 776 Propulsive
MovementsPeristalsis 776 Mixing Movements 777 Gastrointestinal
Blood Flow Splanchnic Circulation 777 Anatomy of the
Gastrointestinal Blood Supply 778 Effect of Gut Activity and
Metabolic Factors on Gastrointestinal Blood Flow 778 Nervous
Control of Gastrointestinal Blood Flow 779 C H A P T E R 6 3
Propulsion and Mixing of Food in the Alimentary Tract 781 Ingestion
of Food 781 Mastication (Chewing) 781 Swallowing (Deglutition) 782
Motor Functions of the Stomach 784 Storage Function of the Stomach
784 Mixing and Propulsion Of Food in the StomachThe Basic
Electrical Rhythm of the Stomach Wall 784 Stomach Emptying 785
Regulation of Stomach Emptying 785 Movements of the Small Intestine
786 Mixing Contractions (Segmentation Contractions) 786 Propulsive
Movements 787 Function of the Ileocecal Valve 788 Movements of the
Colon 788 Defecation 789 Other Autonomic Reexes That Affect Bowel
Activity 790 C H A P T E R 6 4 Secretory Functions of the
Alimentary Tract 791 General Principles of Alimentary Tract
Secretion 791 Anatomical Types of Glands 791 Basic Mechanisms of
Stimulation of the Alimentary Tract Glands 791 Basic Mechanism of
Secretion by Glandular Cells 791 Lubricating and Protective
Properties of Mucus, and Importance of Mucus in the
Gastrointestinal Tract 793 Secretion of Saliva 793 Nervous
Regulation of Salivary Secretion 794 Esophageal Secretion 795
28. xxx Table of Contents Gastric Secretion 794 Characteristics
of the Gastric Secretions 794 Pyloric GlandsSecretion of Mucus and
Gastrin 797 Surface Mucous Cells 797 Stimulation of Gastric Acid
Secretion 797 Regulation of Pepsinogen Secretion 798 Inhibition of
Gastric Secretion by Other Post-Stomach Intestinal Factors 798
Chemical Composition of Gastrin And Other Gastrointestinal Hormones
799 Pancreatic Secretion 799 Pancreatic Digestive Enzymes 799
Secretion of Bicarbonate Ions 800 Regulation of Pancreatic
Secretion 800 Secretion of Bile by the Liver; Functions of the
Biliary Tree 802 Physiologic Anatomy of Biliary Secretion 802
Function of Bile Salts in Fat Digestion and Absorption 804 Liver
Secretion of Cholesterol and Gallstone Formation 804 Secretions of
the Small Intestine 805 Secretion of Mucus by Brunners Glands in
the Duodenum 805 Secretion of Intestinal Digestive Juices by the
Crypts of Lieberkhn 805 Regulation of Small Intestine Secretion
Local Stimuli 806 Secretions of the Large Intestine 806 C H A P T E
R 6 5 Digestion and Absorption in the Gastrointestinal Tract 808
Digestion of the Various Foods by Hydrolysis 808 Digestion of
Carbohydrates 809 Digestion of Proteins 810 Digestion of Fats 811
Basic Principles of Gastrointestinal Absorption 812 Anatomical
Basis of Absorption 812 Absorption in the Small Intestine 813
Absorption of Water 814 Absorption of Ions 814 Absorption of
Nutrients 815 Absorption in the Large Intestine: Formation of Feces
817 C H A P T E R 6 6 Physiology of Gastrointestinal Disorders 819
Disorders of Swallowing and of the Esophagus 819 Disorders of the
Stomach 819 Peptic Ulcer 820 Specic Causes of Peptic Ulcer in the
Human Being 821 Disorders of the Small Intestine 821 Abnormal
Digestion of Food in the Small IntestinePancreatic Failure 821
Malabsorption by the Small Intestine MucosaSprue 822 Disorders of
the Large Intestine 822 Constipation 822 Diarrhea 822 Paralysis of
Defecation in Spinal Cord Injuries 823 General Disorders of the
Gastrointestinal Tract 823 Vomiting 823 Nausea 824 Gastrointestinal
Obstruction 824 U N I T X I I I Metabolism and Temperature
Regulation C H A P T E R 6 7 Metabolism of Carbohydrates, and
Formation of Adenosine Triphosphate 829 Release of Energy from
Foods, and the Concept of Free Energy 829 Role of Adenosine
Triphosphate in Metabolism 829 Central Role of Glucose in
Carbohydrate Metabolism 830 Transport of Glucose Through the Cell
Membrane 831 Insulin Increases Facilitated Diffusion of Glucose 831
Phosphorylation of Glucose 831 Glycogen Is Stored in Liver and
Muscle 831 GlycogenesisThe Process of Glycogen Formation 832
Removal of Stored Glycogen Glycogenolysis 832 Release of Energy
from the Glucose Molecule by the Glycolytic Pathway 832 Summary of
ATP Formation During the Breakdown of Glucose 836 Control of Energy
Release from Stored Glycogen When the Body Needs Addit