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Preface
The first edition of the Textbook of Medical Physiology
was written by Arthur C. Guyton almost 55 years ago.
Unlike most major medical textbooks, which often have
20 or more authors, the first eight editions of the Textbook
of Medical Physiology were written entirely by Dr. Guyton,
with each new edition arriving on schedule for nearly 40
years. The Textbook of Medical Physiology, first published
in 1956, quickly became the best-selling medical physiology textbook in the world. Dr. Guyton had a gift for
communicating complex ideas in a clear and interesting
manner that made studying physiology fun. He wrote the
book to help students learn physiology, not to impress his
professional colleagues.
I worked closely with Dr. Guyton for almost 30 years
and had the privilege of writing parts of the 9th and 10th
editions. After Dr. Guytons tragic death in an automobile
accident in 2003, I assumed responsibility for completing
the 11th edition.
For the 12th edition of the Textbook of Medical
Physiology, I have the same goal as for previous editions
to 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 fun because our
rapidly increasing knowledge of physiology continues to
unravel new mysteries of body functions. Advances in
molecular and cellular physiology have made it possible to explain many physiology principles in the terminology of molecular and physical sciences rather than
in merely a series of separate and unexplained biological
phenomena.
The Textbook of Medical Physiology, however, is not
a reference book that attempts to provide a compendium of the most recent advances in physiology. This is
a book that continues the tradition of being written for
students. It focuses on the basic principles of physiology needed to begin a career in the health care professions, such as medicine, dentistry and nursing, as well
as graduate studies in the biological and health sciences.
It should also be useful to physicians and health care
professionals who wish to review the basic principles
needed for understanding the pathophysiology of
human disease.
I have attempted to maintain the same unified 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 continue to use it during their
professional careers.
My hope is 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 function,
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? Fortunately, our bodies are endowed with a vast network of feedback controls that achieve the necessary balances without which
we would be unable to live. Physiologists call this high
level of internal bodily control homeostasis. In disease
states, functional balances are often seriously disturbed
and homeostasis is impaired. 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 functions in
disease.
Another objective is to be as accurate as possible.
Suggestions and critiques from many students, physiologists, 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 thousands of bits of information, I wish to issue 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 feedback important for
progressive improvement of a textbook of physiology. To
the many persons who have already helped, I express sincere thanks.
vii
Preface
viii
John E. Hall
Contents
UNIT I
40
40
UNIT II
CHAPTER 1
Functional Organization of the Human Body
and Control of the Internal Environment
Cells as the Living Units of the Body
Extracellular FluidThe Internal
Environment
Homeostatic Mechanisms of the Major
Functional Systems
Control Systems of the Body
SummaryAutomaticity of the Body
CHAPTER 2
The Cell and Its Functions
Organization of the Cell
Physical Structure of the Cell
Comparison of the Animal Cell with
Precellular Forms of Life
Functional Systems of the Cell
Locomotion of Cells
CHAPTER 3
Genetic Control of Protein Synthesis, Cell
Function, and Cell Reproduction
Genes in the Cell Nucleus
The DNA Code in the Cell Nucleus Is
Transferred to an RNA Code in the Cell
CytoplasmThe Process of Transcription
Synthesis of Other Substances in the Cell
Control of Gene Function and Biochemical
Activity in Cells
The DNA-Genetic System Also Controls Cell
Reproduction
Cell Differentiation
27
27
30
35
35
37
39
CHAPTER 4
Transport of Substances Through Cell
Membranes
The Lipid Barrier of the Cell Membrane,
and Cell Membrane Transport Proteins
Diffusion
Active Transport of Substances Through
Membranes
CHAPTER 5
Membrane Potentials and Action Potentials
Basic Physics of Membrane Potentials
Measuring the Membrane Potential
Resting Membrane Potential of Nerves
Nerve Action Potential
Roles of Other Ions During the Action
Potential
Propagation of the Action Potential
Re-establishing Sodium and Potassium
Ionic Gradients After Action Potentials Are
CompletedImportance of Energy
Metabolism
Plateau in Some Action Potentials
Rhythmicity of Some Excitable Tissues
Repetitive Discharge
Special Characteristics of Signal Transmission
in Nerve Trunks
ExcitationThe Process of Eliciting the
Action Potential
Recording Membrane Potentials and
Action Potentials
45
45
46
52
57
57
58
59
60
64
64
65
66
66
67
68
69
ix
Contents
CHAPTER 6
Contraction of Skeletal Muscle
Physiologic Anatomy of Skeletal Muscle
General Mechanism of Muscle Contraction
Molecular Mechanism of Muscle Contraction
Energetics of Muscle Contraction
Characteristics of Whole Muscle
Contraction
CHAPTER 7
Excitation of Skeletal Muscle:
Neuromuscular Transmission and
Excitation-Contraction Coupling
Transmission of Impulses from Nerve Endings
to Skeletal Muscle Fibers: The Neuromuscular
Junction
Molecular Biology of Acetylcholine Formation
and Release
Drugs That Enhance or Block Transmission
at the Neuromuscular Junction
Myasthenia Gravis Causes Muscle Paralysis
Muscle Action Potential
Excitation-Contraction Coupling
CHAPTER 11
71
71
73
74
78
79
83
83
86
86
86
87
88
CHAPTER 8
Excitation and Contraction of Smooth Muscle 91
Contraction of Smooth Muscle
91
Nervous and Hormonal Control of Smooth
Muscle Contraction
94
UNIT III
The Heart
CHAPTER 9
Cardiac Muscle; The Heart as a Pump and
Function of the Heart Valves
Physiology of Cardiac Muscle
Cardiac Cycle
Relationship of the Heart Sounds to Heart
Pumping
Work Output of the Heart
Chemical Energy Required for Cardiac Contraction:
Oxygen Utilization by the Heart
Regulation of Heart Pumping
CHAPTER 10
Rhythmical Excitation of the Heart
Specialized Excitatory and Conductive System
of the Heart
Control of Excitation and Conduction in the
Heart
x
101
101
104
107
107
109
110
115
115
118
121
121
123
123
124
129
129
131
134
137
137
138
141
143
143
144
146
148
149
151
152
153
UNIT IV
The Circulation
CHAPTER 14
Overview of the Circulation; Biophysics of
Pressure, Flow, and Resistance
Physical Characteristics of the Circulation
Basic Principles of Circulatory Function
Interrelationships of Pressure, Flow, and
Resistance
157
157
158
159
Contents
CHAPTER 15
Vascular Distensibility and Functions of the
Arterial and Venous Systems
Vascular Distensibility
Arterial Pressure Pulsations
Veins and Their Functions
CHAPTER 16
The Microcirculation and Lymphatic
System: Capillary Fluid Exchange,
Interstitial Fluid, and Lymph Flow
Structure of the Microcirculation
and Capillary System
Flow of Blood in the Capillaries
Vasomotion
Exchange of Water, Nutrients, and Other
Substances Between the Blood and
Interstitial Fluid
Interstitium and Interstitial Fluid
Fluid Filtration Across Capillaries Is
Determined by Hydrostatic and Colloid
Osmotic Pressures, as Well as Capillary
Filtration Coefficient
Lymphatic System
CHAPTER 17
Local and Humoral Control of Tissue
Blood Flow
Local Control of Blood Flow in Response to
Tissue Needs
Mechanisms of Blood Flow Control
Humoral Control of the Circulation
CHAPTER 18
Nervous Regulation of the Circulation,
and Rapid Control of Arterial Pressure
Nervous Regulation of the Circulation
Role of the Nervous System in Rapid
Control of Arterial Pressure
Special Features of Nervous Control
of Arterial Pressure
CHAPTER 19
Role of the Kidneys in Long-Term Control of
Arterial Pressure and in Hypertension: The
Integrated System for Arterial Pressure
Regulation
RenalBody Fluid System for Arterial
Pressure Control
The Renin-Angiotensin System: Its Role
in Arterial Pressure Control
Summary of the Integrated, Multifaceted
System for Arterial Pressure Regulation
167
167
168
171
177
177
178
179
180
181
186
191
191
191
199
201
201
204
209
213
213
220
226
CHAPTER 20
Cardiac Output, Venous Return,
and Their Regulation
Normal Values for Cardiac Output at Rest
and During Activity
Control of Cardiac Output by Venous
ReturnRole of the Frank-Starling Mechanism
of the Heart
Pathologically High or Low Cardiac Outputs
Methods for Measuring Cardiac
Output
CHAPTER 21
Muscle Blood Flow and Cardiac Output
During Exercise; the Coronary Circulation
and Ischemic Heart Disease
Blood Flow Regulation in Skeletal Muscle
at Rest and During Exercise
Coronary Circulation
CHAPTER 22
Cardiac Failure
Circulatory Dynamics in Cardiac Failure
Unilateral Left Heart Failure
Low-Output Cardiac Failure
Cardiogenic Shock
Edema in Patients with Cardiac Failure
Cardiac Reserve
CHAPTER 23
Heart Valves and Heart Sounds;
Valvular and Congenital Heart
Defects
Heart Sounds
Abnormal Circulatory Dynamics in Valvular
Heart Disease
Abnormal Circulatory Dynamics
in Congenital Heart Defects
Use of Extracorporeal Circulation During
Cardiac Surgery
Hypertrophy of the Heart in Valvular
and Congenital Heart Disease
CHAPTER 24
Circulatory Shock and Its Treatment
Physiologic Causes of Shock
Shock Caused by Hypovolemia
Hemorrhagic Shock
Neurogenic ShockIncreased Vascular
Capacity
Anaphylactic Shock and Histamine Shock
Septic Shock
229
229
229
232
240
243
243
246
255
255
259
259
259
261
265
265
268
269
271
272
273
273
274
279
280
280
xi
Contents
280
281
UNIT V
285
285
286
287
287
287
287
289
290
290
291
292
294
294
296
300
303
303
304
307
307
308
309
309
Abnormalities of Micturition
Urine Formation Results from Glomerular
Filtration, Tubular Reabsorption, and Tubular
Secretion
Glomerular FiltrationThe First Step in
Urine Formation
Determinants of the GFR
Renal Blood Flow
Physiologic Control of Glomerular Filtration
and Renal Blood Flow
Autoregulation of GFR and Renal Blood Flow
310
310
312
314
316
317
319
CHAPTER 27
Urine Formation by the Kidneys: II. Tubular
Reabsorption and Secretion
323
Renal Tubular Reabsorption and Secretion
323
Tubular Reabsorption Includes Passive
and Active Mechanisms
323
Reabsorption and Secretion Along Different
Parts of the Nephron
329
Regulation of Tubular Reabsorption
334
Use of Clearance Methods to Quantify Kidney
Function
340
CHAPTER 28
Urine Concentration and Dilution; Regulation
of Extracellular Fluid Osmolarity and Sodium
Concentration
Kidneys Excrete Excess Water by Forming
Dilute Urine
Kidneys Conserve Water by Excreting
Concentrated Urine
Quantifying Renal Urine Concentration
and Dilution: Free Water and Osmolar
Clearances
Disorders of Urinary Concentrating Ability
Control of Extracellular Fluid Osmolarity and
Sodium Concentration
Osmoreceptor-ADH Feedback System
Importance of Thirst in Controlling
Extracellular Fluid Osmolarity and Sodium
Concentration
Salt-Appetite Mechanism for Controlling
Extracellular Fluid Sodium Concentration and
Volume
CHAPTER 29
Renal Regulation of Potassium, Calcium,
Phosphate, and Magnesium; Integration
of Renal Mechanisms for Control of Blood
Volume and Extracellular Fluid Volume
Regulation of Extracellular Fluid Potassium
Concentration and Potassium Excretion
345
345
346
354
354
355
355
357
360
361
361
Contents
367
369
370
371
CHAPTER 31
Diuretics, Kidney Diseases
Diuretics and Their Mechanisms of Action
Kidney Diseases
Acute Renal Failure
Chronic Renal Failure: An Irreversible Decrease
in the Number of Functional Nephrons
Specific Tubular Disorders
Treatment of Renal Failure by Transplantation
or by Dialysis with an Artificial Kidney
397
397
399
399
401
408
409
373
UNIT VI
373
376
376
377
379
379
379
380
380
381
383
383
384
385
386
388
389
391
391
392
393
393
413
413
420
421
423
423
425
426
428
430
431
431
431
433
433
443
445
445
445
447
449
xiii
Contents
CHAPTER 36
Hemostasis and Blood Coagulation
Events in Hemostasis
Vascular Constriction
Mechanism of Blood Coagulation
Conditions That Cause Excessive Bleeding in
Humans
Thromboembolic Conditions in the
Human Being
Anticoagulants for Clinical Use
Blood Coagulation Tests
451
451
451
453
457
459
459
460
UNIT VII
Respiration
CHAPTER 37
Pulmonary Ventilation
465
Mechanics of Pulmonary Ventilation
465
Pulmonary Volumes and Capacities
469
Minute Respiratory Volume Equals Respiratory
Rate Times Tidal Volume
471
Alveolar Ventilation
471
Functions of the Respiratory Passageways
472
CHAPTER 38
Pulmonary Circulation, Pulmonary Edema,
Pleural Fluid
Physiologic Anatomy of the Pulmonary
Circulatory System
Pressures in the Pulmonary System
Blood Volume of the Lungs
Blood Flow Through the Lungs and Its
Distribution
Effect of Hydrostatic Pressure Gradients in
the Lungs on Regional Pulmonary Blood Flow
Pulmonary Capillary Dynamics
Fluid in the Pleural Cavity
CHAPTER 39
Physical Principles of Gas Exchange;
Diffusion of Oxygen and Carbon Dioxide
Through the Respiratory Membrane
Physics of Gas Diffusion and Gas
Partial Pressures
Compositions of Alveolar Air and Atmospheric
Air Are Different
Diffusion of Gases Through the Respiratory
Membrane
Effect of the Ventilation-Perfusion Ratio on
Alveolar Gas Concentration
xiv
477
477
477
478
479
479
481
483
485
485
487
489
492
CHAPTER 40
Transport of Oxygen and Carbon Dioxide in
Blood and Tissue Fluids
Transport of Oxygen from the Lungs to the
Body Tissues
Transport of Carbon Dioxide in the Blood
Respiratory Exchange Ratio
CHAPTER 41
Regulation of Respiration
Respiratory Center
Chemical Control of Respiration
Peripheral Chemoreceptor System for Control
of Respiratory ActivityRole of Oxygen in
Respiratory Control
Regulation of Respiration During Exercise
Other Factors That Affect Respiration
CHAPTER 42
Respiratory InsufficiencyPathophysiology,
Diagnosis, Oxygen Therapy
Useful Methods for Studying Respiratory
Abnormalities
Pathophysiology of Specific Pulmonary
Abnormalities
Hypoxia and Oxygen Therapy
HypercapniaExcess Carbon Dioxide in the
Body Fluids
Artificial Respiration
495
495
502
504
505
505
507
508
510
512
515
515
517
520
522
522
UNIT VIII
527
527
531
533
533
535
535
539
540
540
Contents
UNIT IX
543
543
545
546
546
559
559
560
563
564
564
569
CHAPTER 47
Somatic Sensations: I. General Organization,
the Tactile and Position Senses
571
Classification of Somatic Senses
571
Detection and Transmission of Tactile
Sensations
571
Sensory Pathways for Transmitting Somatic
Signals into the Central Nervous System
573
Transmission in the Dorsal ColumnMedial
Lemniscal System
573
Transmission of Less Critical Sensory Signals
in the Anterolateral Pathway
580
Some Special Aspects of Somatosensory
Function
581
CHAPTER 48
Somatic Sensations: II. Pain, Headache, and
Thermal Sensations
Types of Pain and Their QualitiesFast Pain
and Slow Pain
583
584
586
588
588
590
590
592
UNIT X
557
CHAPTER 46
Sensory Receptors, Neuronal Circuits for
Processing Information
Types of Sensory Receptors and the
Stimuli They Detect
Transduction of Sensory
Stimuli into Nerve Impulses
Nerve Fibers That Transmit Different Types of
Signals and Their Physiologic Classification
Transmission of Signals of Different Intensity
in Nerve TractsSpatial and Temporal
Summation
Transmission and Processing of Signals in
Neuronal Pools
Instability and Stability of Neuronal Circuits
583
583
CHAPTER 49
The Eye: I. Optics of Vision
Physical Principles of Optics
Optics of the Eye
Ophthalmoscope
Fluid System of the EyeIntraocular Fluid
CHAPTER 50
The Eye: II. Receptor and Neural Function
of the Retina
Anatomy and Function of the Structural
Elements of the Retina
Photochemistry of Vision
Color Vision
Neural Function of the Retina
CHAPTER 51
The Eye: III. Central Neurophysiology
of Vision
Visual Pathways
Organization and Function of the Visual
Cortex
Neuronal Patterns of Stimulation During
Analysis of the Visual Image
Fields of Vision; Perimetry
Eye Movements and Their Control
Autonomic Control of Accommodation
and Pupillary Aperture
CHAPTER 52
The Sense of Hearing
Tympanic Membrane and the Ossicular System
Cochlea
Central Auditory Mechanisms
Hearing Abnormalities
597
597
600
605
606
609
609
611
615
616
623
623
624
626
627
627
631
633
633
634
639
642
xv
Contents
CHAPTER 53
The Chemical SensesTaste and Smell
Sense of Taste
Sense of Smell
645
645
648
UNIT XI
703
704
705
CHAPTER 58
655
655
657
661
663
663
663
664
664
665
665
667
667
673
674
678
681
681
689
694
CHAPTER 57
Cerebral Cortex, Intellectual Functions of the
Brain, Learning, and Memory
697
Physiologic Anatomy of the Cerebral Cortex
697
Functions of Specific Cortical Areas
698
xvi
721
721
725
726
727
729
729
731
738
738
739
743
743
746
749
Contents
UNIT XII
Gastrointestinal Physiology
CHAPTER 62
General Principles of Gastrointestinal
FunctionMotility, Nervous Control, and
Blood Circulation
General Principles of Gastrointestinal Motility
Neural Control of Gastrointestinal Function
Enteric Nervous System
Functional Types of Movements in the
Gastrointestinal Tract
Gastrointestinal Blood FlowSplanchnic
Circulation
CHAPTER 63
Propulsion and Mixing of Food in the
Alimentary Tract
Ingestion of Food
Motor Functions of the Stomach
Movements of the Small Intestine
Movements of the Colon
Other Autonomic Reflexes That Affect Bowel
Activity
CHAPTER 64
Secretory Functions of the Alimentary Tract
General Principles of Alimentary Tract
Secretion
Secretion of Saliva
Esophageal Secretion
Gastric Secretion
Pancreatic Secretion
Secretion of Bile by the Liver; Functions of the
Biliary Tree
Secretions of the Small Intestine
Secretion of Mucus by the Large Intestine
CHAPTER 65
Digestion and Absorption in the
Gastrointestinal Tract
Digestion of the Various Foods by Hydrolysis
Basic Principles of Gastrointestinal Absorption
Absorption in the Small Intestine
Absorption in the Large Intestine: Formation of
Feces
CHAPTER 66
Physiology of Gastrointestinal Disorders
Disorders of Swallowing and of the Esophagus
753
753
755
759
759
763
763
765
768
770
772
773
773
775
776
777
780
783
786
787
789
789
793
794
797
799
799
799
801
802
803
UNIT XIII
809
810
810
811
812
816
817
817
819
819
821
822
825
826
827
831
831
831
833
835
837
837
837
839
840
xvii
Contents
CHAPTER 71
Dietary Balances; Regulation of Feeding;
Obesity and Starvation; Vitamins and
Minerals
843
Energy Intake and Output Are Balanced Under
Steady-State Conditions
843
Dietary Balances
843
Regulation of Food Intake and Energy
Storage
845
Obesity
850
Inanition, Anorexia, and Cachexia
851
Starvation
852
Vitamins
852
Mineral Metabolism
855
CHAPTER 72
Energetics and Metabolic Rate
Adenosine Triphosphate (ATP) Functions as
an Energy Currency in Metabolism
Control of Energy Release in the Cell
Metabolic Rate
Energy MetabolismFactors That Influence
Energy Output
CHAPTER 73
Body Temperature Regulation,
and Fever
Normal Body Temperatures
Body Temperature Is Controlled by
Balancing Heat Production and
Heat Loss
Regulation of Body Temperature
Role of the Hypothalamus
Abnormalities of Body Temperature
Regulation
859
859
861
862
863
867
867
867
871
875
UNIT XIV
881
881
881
884
886
891
CHAPTER 75
Pituitary Hormones and Their Control by the
Hypothalamus
Pituitary Gland and Its Relation to the
Hypothalamus
Hypothalamus Controls Pituitary Secretion
Physiological Functions of Growth Hormone
Posterior Pituitary Gland and Its Relation to
the Hypothalamus
CHAPTER 76
Thyroid Metabolic Hormones
Synthesis and Secretion of the Thyroid
Metabolic Hormones
Physiological Functions of the Thyroid
Hormones
Regulation of Thyroid Hormone Secretion
Diseases of the Thyroid
CHAPTER 77
Adrenocortical Hormones
Synthesis and Secretion of Adrenocortical
Hormones
Functions of the Mineralocorticoids
Aldosterone
Functions of the Glucocorticoids
Adrenal Androgens
Abnormalities of Adrenocortical Secretion
CHAPTER 78
Insulin, Glucagon, and Diabetes Mellitus
Insulin and Its Metabolic Effects
Glucagon and Its Functions
Somatostatin Inhibits Glucagon and Insulin
Secretion
Summary of Blood Glucose Regulation
Diabetes Mellitus
CHAPTER 79
Parathyroid Hormone, Calcitonin, Calcium
and Phosphate Metabolism, Vitamin D, Bone,
and Teeth
Overview of Calcium and
Phosphate Regulation in the Extracellular
Fluid and Plasma
Bone and Its Relation to Extracellular Calcium
and Phosphate
Vitamin D
Parathyroid Hormone
Calcitonin
Summary of Control of Calcium Ion
Concentration
895
895
897
898
904
907
907
910
914
916
921
921
924
928
934
934
939
939
947
949
949
950
955
955
957
960
962
966
966