textbook of medical physiology€¦ · chapter 2 the cell and its function 12 organization of the...
TRANSCRIPT
TEXTBOOK OF
MEDICALPHYSIOLOGY
SIXTH EDITION
ARTHUR C. GUYTON, M.D.Chairman and Professor of the Department of Physiology
and Biophysics, University of Mississippi, School of Medicine
FACHBEREICH BIO'.OGIE (10)der Technischec '•• tadi:;chuta-Darmstadt
/ t . ; •• i • , • . i .
— b I 3 I ! • ' 1 i i J !v —
D - 6100 Do r m s i ; ; d t / B. R. D.
SchniHsoohnstrafie'
Inv.-Nr.__!
1981 W. B. SAUNDERS COMPANY Philadelphia London Toronto
CONTENTS
PART IINTRODUCTION TO PHYSIOLOGY:
THE CELL AND GENERAL PHYSIOLOGY
CHAPTER 1
FUNCTIONAL ORGANIZATION OFTHE HUMAN BODY AND CONTROLOF THE "INTERNALENVIRONMENT" 2Cells as the Living Units of the Body 2The Extracellular Fluid-The Internal
Environment 2"Homeostatic" Mechanisms*of the
Major Functional Systems 3Homeostasis 3The Extracellular Fluid Transport
System 3Origin of Nutrients in the
Extracellular Fluid 4Removal of Metabolic End-
Products 4Regulation of Body Functions 4Reproduction 5
The Control Systems of the Body '. 5Examples of Control Mechanisms... 5Characteristics of Control Systems... 6Automaticity of the Body , 7Appendix: Basic Physical
Principles of Control Systems 7Oscillation of Control Systems 10
CHAPTER 2
THE CELL AND ITS FUNCTION 12Organization of the Cell 12Physical Structure of the Cell 13
The Membranous Structures ofthe Cell 13
The Cytoplasm and Its Organelles... 16The Nucleus 17
Comparison of the Animal Cell withPrecellular Forms of Life 18
Functional Systems of the Cell 19Ingestion by the Cell 19Digestion of Foreign Substances
in the Cell — Function of theLysosomes 20
Extraction of Energy fromNutrients — Function of theMitochondria 21
Synthesis and Formation of" Cellular Structures by theEndoplasmic Reticulum andthe Golgi Complex 23
Cell Movement 24
CHAPTER 3
GENETIC CONTROL OF CELLFUNCTION-PROTEIN SYNTHESISAND CELL REPRODUCTION 28
The Genes 28The Genetic Code 30
Ribonucleic Acid (RNA) - TheProcess of Transcription 30
Messenger RNA 31Transfer RNA 31Ribosomal RNA 32Formation of Proteins in the
Ribosomes —The Processof Translation 32
Synthesis of Other Substances inthe Cell 34
Control of Genetic Function and Bio-chemical Activity in Cells 34
Genetic Regulation 34Control of Enzyme Activity 35
VIII CONTENTS
Cell Reproduction 36Replication of the DNA 36The Chromosomes and Their
Replication 36Mitosis -. 36Control of Cell Growth and
Reproduction 37Cell Differentiation " 38Cancer...., 38
CHAPTER 4
TRANSPORT THROUGH THE CELLMEMBRANE 41Diffusion 41
Kinetics of Diffusion —TheConcentration Difference 42
Diffusion Through the CellMembrane 42
Active Transport 50Basic Mechanism of Active
Transport 50Transport Through Intracellular
Membranes 51Active Transport Through
Cellular Sheets 51Secondary Active Transport:
Sodium of "Co-Transport"Glucose and Amino Acids 51
Active Transport of Specific Substances... 52Active Transport of Sodium and
Potassium 52Active Transport of Other Ions 53Transport of Sugars 53Transport of Amino Acids 53Pinocytosis and Phagocytosis 54
PART IIBLOOD CELLS, IMMUNITY, AND BLOOD CLOTTING
CHAPTER 5
RED BLOOD CELLS, ANEMIA,AND POLYCYTHEMIA 56The Red Blood Cells 56
Production of Red Blood Cells 57Formation of Hemoglobin 60Iron Metabolism 61
Destruction of Red Blood Cells 62The Anemias 62
Effects of Anemia on theCirculatory System 63
Polycythemia 63Effect of Polycythemia on the
Circulatory System 64
CHAPTER*^
RESISTANCE OF THE BODY TOINFECTION-THE LEUKOCYTES,THE TISSUE MACROPHAGE SYSTEM,AND INFLAMMATION 65The Leukocytes (White Blood Cells) 65
General Characteristics ofLeukocytes 65
Genesis of the Leukocytes 66
Life Span of the White BloodCells 66
Properties of Neutrophils,Monocytes, and Macrophages 67
The Tissue Macrophage System(The ReticuloendothelialSystem) 68
Inflammation and Function ofNeutrophils and Macrophages 70
The Process of Inflammation 70Neutrophilia Caused by Condi-
tions Other Than Inflammation ... 71The Eosinophils 71The Basophils 72
Agranulocytosis 72The Leukemias 72
Effects of Leukemia on theBody 73
CHAPTER 7
IMMUNITY AND ALLERGY 74Innate Immunity 74
Acquired Immunity (or AdaptiveImmunity) 74
CONTENTS
Two Basic Types of AcquiredImmunity 74
Antigens 75Role of Lymphoid Tissue in
Acquired Immunity 75Preprocessing of the T and B'
Lymphocytes 76Mechanisms for Determining
Specificity of SensitizedLymphocytes and Anti-Bodies—Lymphocyte Clones 76
Tolerance of the AcquiredImmunity System to One's OwnTissues —Role of the Thymusand the Bursa 77
Specific Attributes of the BLymphocyte System— "Humoral Immunity and theAntibodies 77
Special Attributes of the TLymphocyte System —Cellular Immunity andSensitized Lymphocytes , -80
Vaccination 81Passive Immunity 81
Interferon — Another Type ofAcquired Immunity 81
Allergy , 82Allergies That Occur in Normal
People .->. 82Allergies in the "Allergic"
Person 82
CHAPTER 8
BLOOD GROUPS; TRANSFUSION;TISSUE AND ORGAN TRANS-PLANTATION 84
Antigenicity and ImmuneReactions of Blood ': 84
0-A-B Blood Groups -.'.... 84The A and B Antigens —Called
"Agglutinogens" 84The Agglutinins 85The Agglutination Process in
Transfusion Reactions 85Blood Typing 86Cross-Matching 86
The Rh Blood Types 86The Rh Immune Response 87
Other Blood Factors 88Transfusion 88
Transfusion Reactions Resultingfrom Mismatched Blood Groups... 88
Other Types of TransfusionReactions 89
Transplantation of Tissues and Organs.Attempts to Overcome the
Antigen-Antibody Reactionsin Transplanted Tissue
IX
89
90
CHAPTER 9
HEMOSTASIS AND BLOODCOAGULATION 92Events in Hemostasis 92
Vascular Spasm 92Formation of the Platelet Plug 92Blood Coagulation in the
Ruptured Vessel 93Fibrous Organization or
Dissolution of the Blood Clot 93Mechanism of Blood Coagulation 93
Conversion of Prothrombin toThrombin 93
Conversion of Fibrinogen toFibrin — Formation of the Clot 94
The Vicious Cycle of Clot .Formation 94
Block of Clot Growth by BloodFlow 95
Initiation of Coagulation: Forma-tion of Prothrombin Activator 95
Prevention of Blood Clotting inthe Normal Vascular System —The IntravascularAnticoagulants 98
Lysis of Blood Clots —Plasmin 98Conditions that Cause Excessive
Bleeding in Human Beings 99Decreased Prothrombin, Factor
VII, Factor IX, and Factor XCaused by Vitamin K Deficiency 99
Hemophilia 99Thrombocytopenia 99
Thromboembolic Conditions in theHuman Being 100
Femoral Thrombosis and Mas-sive Pulmonary Embolism 100
Disseminated IntravascularClotting 100
Anticoagulants for Clinical Use 100Heparin as an Intravenous
Anticoagulant 100Coumarins as Anticoagulants 101Prevention of Blood Coagulation
Outside the Body 101Blood Coagulation Tests 101
Bleeding Time 101Clotting Time 101Prothrombin Time 101
CONTENTS
PART IIINERVE AND MUSCLE
CHAPTER 10
MEMBRANE POTENTIALS, ACTIONPOTENTIALS, EXCITATION, ANDRHYTHMICITY 104Basic Physics of Membrane Potentials 104
Membrane Potentials Caused byActive Transport—The"Electrogenic Pump" 104
Membrane Potentials Caused byDiffusion 105
Calculation of the Membrane PotentialWhen the Membrane Is Permeableto Several Different Ions 106
Origin of the Nerve Cell MembranePotential : 106
The Action Potential 109Relationship of the Action
Potential to the Potassium andSodium Nernst Potentials....:..".... I l l
Some Experimental MethodsThat Have Been Used to Studythe Action Potential 112
Propagation of the ActionPotential ..:.... 112
"Recharging" the FiberMembrane —Importance of a
Energy Metabolism 113The Spike Potential and the
After-Potentials 114Plateau in the Action Potential 114Rhythmicity of Certain Excitable
Tissues — Repetitive Discharge..... 115Special Aspects of Impulse
Transmission in Nerves 116Velocity of Conduction in Nerve
Fibers 117Excitation—The Process of Eliciting
the Action Potential 117Factors That Increase Membrane
Excitability 119Inhibition of Excitability —
"Stabilizers" and LocalAnesthetics 119
Recording Membrane Potentials andAction Potentials 119
CHAPTER 11
- CONTRACTION OF SKELETAL•MUSCLE 122Physiologic Anatomy of Skeletal
Muscle 122
The Skeletal Muscle Fiber 122Molecular Mechanism of Muscle
Contraction 124Molecular Characteristics of the
Contractile Filaments 125Relationship Between Actin and
Myosin Filament Overlap andTension Developed by theContracting Muscle 127
Relation of Velocity of Contrac-tion to Load :.. 128
Initiation of Muscle Contraction:Excitation-Contraction Coupling 129
The Muscle Action Potential 129Spread of the Action Potential to
the Interior of the Muscle Fiberby Way of the TransverseTubule System 129
Release of Calcium Ions by theCisternae of the SarcoplasmicReticulum 130
The Source of Energy for MuscleContraction 131
Characteristics of a Single MuscleTwitch 132
Mechanics of Skeletal MuscleContraction 133
The Motor Unit 133Summation of Muscle
Contraction 133Skeletal Muscle Tone 134Muscle Fatigue 135The Lever Systems of the Body 135
Special Features and Abnormalities ofSkeletal Muscle Function 135
Muscle Hypertrophy 135Muscle Atrophy 136Rigor Mortis 136Familial Periodic Paralysis 136The Electromyogram 136
CHAPTER 12
NEUROMUSCULAR TRANSMISSION;FUNCTION OF SMOOTH MUSCLE 138Transmission of Impulses from Nerves
to Skeletal Muscle Fibers: TheNeuromuscular Junction 138
Myasthenia Gravis 140Contraction of Smooth Muscle 140
Types of Smooth Muscle 141The Contractile Process in
Smooth Muscle 141
CONTENTS XI
Membrane Potentials and ActionPotentials in Smooth Muscle 142
Excitation-Contraction Coupling —Role of Calcium Ions 144
Neuromuscular Junctions ofSmooth Muscle 145
Smooth Muscle ContractionWithout Action Potentials —Effect of Local Tissue Factorsand Hormones
Mechanical Characteristics ofSmooth Muscle Contraction...
146
146
PART IVTHE HEART
CHAPTER 13
HEART MUSCLE; THE HEART ASA PUMP 150Physiology of Cardiac Muscle .o... 150
Physiologic Anatomy of CardiacMuscle 150
Action Potentials in CardiacMuscle 151
Contraction of Cardiac Muscle ....... 152The Cardiac Cycle 153
Systole and Diastole .'. 153Relationship of the Electrocardio-
gram to the Cardiac Cycle 154Function of the Atria as Pumps 154Function of the Ventricles as
Pumps 155Function of the Valves x 156The Aortic Pressure Curve 156Relationship of the Heart Sounds
to Heart Pumping 157Work Output of the Heart 157Energy for Cardiac Contraction 157
Regulation of Cardiac Function 158Intrinsic Autoregulation of
Cardiac Pumping —The Frank-Starling Law of the Heart 158
Control of the Heart by Nerves 159Effect of Heart Debility on Cardiac
Function — The HypoeffectiveHeart 160
Effect of Exercise on the Heart—The Hypereffective Heart 161
Effect of Various Ions on HeartFunction 161
Effect of Temperature on theHeart 162
The Heart-Lung Preparation 162Assessment of Contractility 163
CHAPTER 14
RHYTHMIC EXCITATION OF THEHEART 165The Special Excitatory and Conductive
System of the Heart 165The Sino-Atrial Node 165
Internodal Pathways and Trans-mission of the Cardiac ImpulseThrough the Atria 166
The Atrioventricular (A-V) Nodeand the Purkinje System 166
Transmission in the PurkinjeSystem 167
Transmission of the CardiacImpulse in the VentricularMuscle 168
Summary of the Spread of theCardiac Impulse Through theHeart '. 168
Control of Excitation and Conductionin the Heart 168
The S-A Node as the Pacemakerof the Heart 168
Role of the Purkinje System inCausing Synchronous Con-traction of the VentricularMuscle 169
Function of the Purkinje Systemin Preventing Arrhythmias 169
Control of Heart Rhythmicityand Conduction by theAutonomic Nerves 170
Abnormal Rhythms of the Heart 171Premature Contractions — Ectopic
Foci.. 171Heart Block 171Flutter and Fibrillation 171Cardiac Arrest 175
CHAPTER 15
THE NORMALELECTROCARDIOGRAM 176Characteristics of the Normal
Electrocardiogram 176Depolarization Waves Versus
Repolarization Waves 176Relationship of Atrial and Ven-
tricular Contraction to the Wavesof the Electrocardiogram 177
Voltage and Time Calibration ofthe Electrocardiogram 177
•v
XII CONTENTS
Methods for RecordingElectrocardiograms 178
The Pen Recorder 178Recording Electrocardiograms
with the Oscilloscope 178Flow of Current Around the Heart
During the Cardiac Cycle 178Recording Electrical Potentials
from a Partially DepolarizedMass of Syncytial CardiacMuscle 178
Flow of Electrical CurrentsAround the Heart in the Chest 179
Electrocardiographic Leads 179The Three Standard Limb Leads 179Chest Leads (Precordial Leads) 181Augmented Unipolar Limb Leads ... 181
CHAPTER 16 .
ELECTROCARDIOGRAPHICINTERPRETATION IN CARDIACMYOPATHIES - VECTORIALANALYSIS.. 182Principles of Vectorial Analysis of >
Electrocardiograms 182Use of Vectors to Represent
Electrical Potentials... 182Denoting the Direction of a Vector
in Terms of Degrees 183"Axis" of Each of the Standard
and Unipolar Leads 183Vectorial Analysis of Potentials
Recorded in Different Leads 183Vectorial Analysis of the Normal
Electrocardiogram 184Vectors Occurring During De-
polarization of the Ventricles —The QRS Complex 184
The Electrocardiogram DuringRepolarization-The T Wave 186
Depolarization of the Atria —Ther 'PWave 186'The Vectorcardiogram 187
The Mean Electrical Axis of theVentricle 187
Determining the Electrical Axisfrom Standard LeadElectrocardiograms 188
Abnormal Ventricular ConditionsThat Cause Axis Deviation 188
Conditions That Cause AbnormalVoltages of the QRS Complex 190
•* Increased Voltage in theStandard Leads 190
Decreased Voltage in theStandard Leads 191
Prolonged and Bizarre Patterns of theQRS Complex 191
Prolonged QRS Complex as aResult of Cardiac Hypertrophyor Dilatation 191
Prolonged QRS Complex Result-ing from Purkinje SystemBlocks 191
Conditions Causing Bizarre QRSComplexes 191
Current of Injury 192Effect of Current of Injury on the
QRS Complex 192The J Point—The Zero Reference
Potential of the Electro-cardiogram 192
Coronary Ischemia as a Cause ofCurrent of Injury 193
Abnormalities in the T Wave 195Effect of Slow Conduction of the
Depolarization Wave on theT Wave 195
Prolonged Depolarization inPortions of the VentricularMuscle as a Cause of Abnor-malities in the T Wave 195
CHAPTER 17 • .
ELECTROCARDIOGRAPHICINTERPRETATION OF CARDIACARRHYTHMIAS 197Abnormal Sinus Rhythms 197
Tachycardia 197Bradycardia 197Sinus Arrhythmia 198
Abnormal Rhythms Resulting fromImpulse Conduction Block 198
Sino-Atrial Block 198Atrioventricular Block 198Incomplete Intraventricular
Block—Electrical Alterhans 199Premature Contractions 200
Premature Atrial Contractions 200A-V Nodal or A-V Bundle
Premature Contractions 200Premature Ventricular Con-
tractions (PVCs) 200Paroxysmal Tachycardia 201
Atrial Paroxysmal Tachycardia 201Ventricular Paroxysmal
Tachycardia 202Abnormal Rhythms Resulting from
Circus Movements 202Atrial Flutter : 202Atrial Fibrillation 202Ventricular Fibrillation 202
CONTENTS XIII
PART VTHE CIRCULATION
CHAPTER 18 t
PHYSICS OF BLOOD, BLOODFLOW, AND PRESSURE:HEMODYNAMICS 206
The Circulatory System as a"Circuit" . 206
The Physical Characteristics of Blood 206The Hematocrit ' 206Plasma 208
Interrelationships Among Pressure,Flow, and Resistance 208
Blood Flow 208Blood Pressure , 210Resistance of Blood Flow ..., 211
Vascular Distensibility — Pressure-Volume Curves 214
Vascular Compliance (orCapacitance) 215
Volume-Pressure Curves of theArterial and VenousCirculations 215
"Mean Circulatory Filling Pressure" andVolume-Pressure Curves of the EntireCirculatory System 216
The Mean Circulatory FillingPressure 216
Volume-Pressure Curves of theEntire Circulation 216
Relationship Between MeanCirculatory Filling Pressure,"Mean Systemic FillingPressure," and "MeanPulmonary Filling Pressure".. 216
Delayed Compliance (Stress-Relaxation)of Vessels 217
CHAPTER 19
THE SYSTEMIC CIRCULATION 219Physical Characteristics of the
Systemic Circulation....; 219Pressure Pulses in the Arteries 221
Factors That Affect the PulsePressure 221
Abnormal Pressure PulseContours 222
Transmission of the PressurePulse to the Periphery 222
The Radial Pulse 224
The Arterioles and Capillaries :. 224Exchange of Fluid Through the
Capillary Membrane 224The Veins and Their Functions 225
Right Atrial Pressure (CentralVenous Pressure) and ItsRelation to Venous Pressure 225
Venous Resistance andPeripheral Venous Pressure 226
Effect of Hydrostatic Pressureon Venous Pressure 226
Venous Valves and the "VenousPump" 227
Pressure Reference Level forMeasuring Venous and OtherCirculatory Pressures 228
Pressure Pulses in the Veins 229Measurement of Venous Pressure ... 229Blood Reservoir Function of the
Veins 229Assessment of Venous Function
by Measurement of MeanCirculatory Filling Pressure 230
CHAPTER 20
LOCAL CONTROL OF BLOOD FLOWBY THE TISSUES, AND NERVOUSAND HUMORAL REGULATION 232Local Control of Blood Flow by the
Tissues Themselves 232Special Types of Local Blood
Flow Regulation 236Long-Term Local Blood Flow
Regulation 237Development of Collateral
Circulation as a Phenomenonof Long-Term Local Blood FlowRegulation : 238
Significance of Long-Term LocalRegulation—The MetabolicMass to Tissue VascularityProportionality 238
Nervous Regulation of the Circulation...... 238The Autonomic Nervous System 238"Patterns" of Circulatory
Responses Elicited by DifferentCentral Nervous SystemCenters 242
Reflex Regulation of theCirculation 242
Humoral Regulation of the Circulation 243
XIV CONTENTS
CHAPTER 21
SHORT-TERM REGULATION OFMEAN ARTERIAL PRESSURE:NERVOUS REFLEX AND HORMONALMECHANISMS FOR RAPID PRESSURE"CONTROL 246Normal Arterial Pressures 246
' The Mean Arterial Pressure 247Clinical Methods for Measuring
Systolic and DiastolicPressures 247
Relationship of Arterial Pressure toCardiac Output and TotalPeripheral Resistance 248
The Overall System for ArterialPressure Regulation 248
Rapidly Acting Nervous Mechanismsfor Arterial Pressure Control 249
The Arterial BaroreceptorControl System —BaroreceptorReflexes 249
Control of Arterial Pressure by -the Carotid and AorticChemoreceptors — Effect ofOxygen Lack on Arterial 'Pressure '. '.'*. 252
Atrial and Pulmonary ArteryReflexes That Help toRegulate Arterial Pressure 252
Control of Arterial Pressure bythe Vasomotor Center inResponse to Diminished BrainBlood Flow-The CNS IschemicResponse. 253
Participation of the Veins inNervous Regulation of CardiacOutput and Arterial Pressure 254
Role of the Skeletal Nerves andSkeletal Muscles to IncreaseCardiac Output and ArterialPressure 254
Respiratory Waves in the ArterialPressure 255
Arterial Pressure VasomotorWaves —Oscillation of the Pres-sure Reflex Control Systems 255
Hormonal Mechanisms for Rapid Controlof Arterial Pressure 255
The Norepinephrine-EpinephrineVasoconstrictor Mechanism 255
The Renin-Angiotensin Vasocon-strictor Mechanism for Controlof Arterial Pressure 256
Role of Vasopressin in RapidControl of Arterial Pressure 257
Two Intrinsic Circulatory Mechanismsfor Arterial Pressure Regulation 257
CHAPTER 22
LONG-TERM REGULATION OF MEANARTERIAL PRESSURE: THE RENAL-BODY FLUID PRESSURE CONTROLSYSTEM; LONG-TERM FUNCTIONSOF THE RENIN-ANGIOTENSINSYSTEM; AND MECHANISMS OFHYPERTENSION 259Short-Term Versus Long-Term Pressure
Control Mechanisms 259The Renal-Body Fluid System for
Arterial Pressure Control 259Factors That Increase the Effec-
tiveness of the Renal-BodyFluid System of PressureControl: (1) The Renin-Angiotensin System, (2) TheAldosterone System, and (3)The Nervous System 261
Graphical Analysis of theFunction of the Kidneys inArterial Pressure Control 262
Respective Roles of TotalPeripheral Resistance andCardiac Output in the Long-Term Control of ArterialPressure 263
Role of the Renin-Angiotensin-Aldosterone System in Long-TermControl of Arterial Pressure 264
Hypertension (High Blood Pressure) 265Some Characteristic Types of
Hypertension 265Volume-Loading Hypertension 265Vasoconstrictor Hypertension —
Hypertension Caused by Con-tinuous Infusion of AngiotensinII or by a Renin-SecretingTumor 267
Goldblatt Hypertension 2 69Hypertension in Toxemia of
Pregnancy 270Neurogenic Hypertension 270Hypertension Caused by
Primary Aldosteronism 270Essential Hypertension 270
Types of Kidney Disease That CauseHypertension; Types That CauseUremia 271
Effects of Hypertension on theBody , 272
CHAPTER 23
CARDIAC OUTPUT, VENOUSRETURN, AND THEIR REGULATION ... 274
Normal Values for CardiacOutput 274
CONTENTS xv
Regulation of Cardiac OutputPrimary Role of the Peripheral
Circulation in the Control ofCardiac Output; Permissive Roleof the Heart :
Role of Total Peripheral Resistancein Determining Normal VenousReturn and Cardiac Output
Effect of Arteriovenous Fistulaeon Cardiac Output
Importance of the "Mean SystemicFilling Pressure" in CardiacOutput Regulation
Regulation of Cardiac Output inHeavy Exercise, Requiring ^Simultaneous Peripheral andCardiac Adjustments :
Abnormally Low and Abnormally HighCardiac Outputs
Low OutputHigh Cardiac Output—Role of
Reduced Total PeripheralResistance in Chronic HighCardiac Output Conditions
Graphical Analysis of Cardiac OutputRegulation y.
Cardiac Output CurvesVenous Return CurvesAnalysis of Cardiac Output and
Right Atrial Pressure UsingCardiac Output and VenousReturn Curves
Right Ventricular Output Versus LeftVentricular Output —BalanceBetween the Ventricles
Methods for Measuring Cardiac OutputPulsatile Output of the Heart as
Measured by an Electromagneticor Ultrasonic Flowmeter.:
Measurement of Cardiac Outputby the Oxygen Fick Method
The Indicator Dilution Method.
278
278
279279
280
281281282
284
286286
286
286287
CHAPTER 24
THE PULMONARY CIRCULATION, , 289Physiologic Anatomy of the Pulmonary
Circulatory System, Pressures in the Pulmonary System
The Blood Volume of the LungsBlood Flow Through the Lungs
; and Its Distribution\ Effect of Hydrostatic Pressurejj Gradients in the Lungs on Regionalf Pulmonary Blood FlowI Effect of Increased Cardiac Output
on the Pulmonary CirculationDuring Heavy Exercise.:
289290291
291
292
293
275 Function of the Pulmonary Circu-lation When the Left AtrialPressure Rises as a Result ofLeft Heart Failure
275 Pulmonary Capillary DynamicsCapillary Exchange of Fluid in
the Lungs276 Pathological Conditions That
Obstruct Blood Flow Through277 the Lungs
293294
294
295
CHAPTER 25
THE CORONARY CIRCULATION ANDISCHEMIC HEART DISEASENormal Coronary Blood Flow and Its
VariationsPhysiologic Anatomy of the
Coronary Blood SupplyNormal Coronary Blood Flow
Control of Coronary Blood FlowLocal Metabolism as the Primary
Controller of Coronary FlowNervous Control of Coronary
Blood FlowThe Substrates of Cardiac
MetabolismIschemic Heart Disease
Myocardial InfarctionCauses of Death Following Acute
Coronary OcclusionThe Stages of Recovery from
Acute Myocardial InfarctionFunction of the Heart Following
Recovery from MyocardialInfarction
Pain in Coronary DiseaseAngina PectorisSurgical Treatment of Coronary
DiseaseMeasurement of Coronary Blood
Flow in Man
298
298
298298299
299
301
302302303
304
305
306306306
306
307
CHAPTER 26
CARDIAC FAILUREDynamics of the Circulation in
Cardiac FailureAcute Effects of Moderate Cardiac
FailureThe Chronic Stage of Failure ........Summary of the Changes That
Occur Following Acute CardiacFailure —"Compensated HeartFailure " .*
309
309
309310
311
XVI CONTENTS
Dynamics of Severe CardiacFailure —Decompensated HeartFailure 312
Unilateral Cardiac Failure .- 313Unilateral Left Heart Failure ~. - 313Unilateral Right Heart Failure. 314
"High Cardiac Output Failure" —Overloading of the Heart 314
Cardiogenic Shock 315Edema in Patients with Cardiac
Failure 315Physiological Classification of Cardiac
Failure 316Cardiac Reserve 317Appendix to Chapter 26: A Quantitative
Graphical Method for Analysis ofCardiac Failure 317
CHAPTER 27
HEART SOUNDS; DYNAMICS OFVALVULAR AND CONGENITALHEART DEFECTS 321The Heart Sounds :..„ 321
Normal Heart Sounds T. :. 321Areas for Auscultation of Normal
Heart Sounds 322The Phonocardiogram ;..'. 323Valvular Lesions 323Abnormal Heart Sounds Caused
by Valvular Lesions 324Abnormal Circulatory Dynamics in
Valvular Heart Disease 325Dynamics of the Circulation in
Aortic Stenosis,and AorticRegurgitation 325
Dynamics of Mitral Stenosis andMitral Regurgitation.~ 326
Circulatory Dynamics DuringExercise in Patients withValvular Lesions 326
Abnormal Circulatory Dynamics inCongenital Heart Defects 326
Patent Ductus Arteriosus — A Left-to-Right Shunt 327
Interventricular Septal Defect —A Left-to-Right Shunt 328
Interatrial Septal Defects — ALeft-to-Right Shunt 329
Tetralogy of Fallot-A Right-to-Left Shunt 329
Pulmonary Stenosis 330• Use of Extracorporeal Circulation
During Cardiac Surgery ^ 330Hypertrophy of the Heart in Valvular
and Congenital Heart Disease 330
CHAPTER 28
CIRCULATORY SHOCK AND PHYSI-OLOGY OF ITS TREATMENT 332
Physiological Causes of Shock ..< 332Shock Caused by Hypovolemia—
Hemorrhagic Shock 332Relationship of Bleeding Volume
to Cardiac Output and ArterialPressure 333
Nonprogressive and ProgressiveHemorrhagic Shock 333
Irreversible Shock 338Hypovolemic Shock Caused by
Plasma Loss 339Hypovolemic Shock Caused by '
Trauma 339Neurogenic Shock —Increased Vascular
Capacity 339Anaphylactic Shock 340Septic Shock 340Effects of Shock on the Body 341Physiology of Treatment in Shock 341
Replacement Therapy 341Treatment of Shock with
Sympathomimetic and Sym-patholytic Drugs or OtherTherapy 342
Other Therapy : 342Circulatory Arrest , 342
Effect of Circulatory Arrest on theBrain 342
CHAPTER 29
MUSCLE BLOOD FLOW DURINGEXERCISE; CEREBRAL, SPLANCHNIC,AND SKIN BLOOD FLOWS 344Blood Flow Through Skeletal Muscles
and Its Regulation in Exercise 344Rate of Blood Flow Through the
Muscles 344Control of Blood Flow Through
the Skeletal Muscles 345Circulatory Readjustments During
Exercise 345The Cerebral Circulation 347
Normal Rate of Cerebral BloodFlow 347
Regulation of Cerebral BloodFlow 347
The Splanchnic Circulation 349Blood Flow Through the Liver 349Blood Flow Through the Intestinal
Vessels 350Portal Venous Pressure 351The Splenic Circulation 351
CONTENTS XVII
Circulation in the Skin 352Physiologic Anatomy of the-
Cutaneous Circulation 352Regulation of Blood Flow in the
Skin 353Color of the Skin in Relation to
Skin Temperature 354
Physiology of Vascular Diseases of the. Limbs 354
Raynaud's Disease 354Buerger's Disease •••• 355Peripheral Arteriosclerosis 355
PART VITHE BODY FLUIDS AND KIDNEYS
CHAPTER 30 "
CAPILLARY DYNAMICS, ANDEXCHANGE OF FLUID BETWEENTHE BLOOD AND INTERSTITIALFLUID.. .: 358
Structure of the Capillary System... 358Flow of Blood in the Capillaries —
Vasomotion '. 359Average Function of the Capillary
System -..-. :.... 359Exchange of Nutrients and Other
Substances Between the Blood andInterstitial Fluid 359
Diffusion Through the CapillaryMembrane 359
The Interstitium and the InterstitialFluid.. 362
Distribution of Fluid Volume Betweenthe Plasma and Interstitial Fluid 363
Capillary Pressure 363Interstitial Fluid Pressure —
Intragel Pressure and FreeFluid Pressure 364
Plasma Colloid Osmotic Pressure ... 366Interstitial Fluid Colloid Osmotic
Pressure 367Exchange of Fluid Volume
Through the CapillaryMembrane 367
The Starling Equilibrium forCapillary Exchange 368
CHAPTER 31
THE LYMPHATIC SYSTEM, INTER-STITIAL FLUID DYNAMICS, EDEMA,AND PULMONARY FLUID v 370The Lymphatic System 370
The Lymph Channels of theBody 370
Formation of Lymph 371Total Rate of Lymph Flow 371
Control of Interstitial Fluid ProteinConcentration and Interstitial FluidPressure 373
Regulation of Interstitial FluidProtein by Lymphatic Pumping... 373
Mechanism of Negative InterstitialFluid Pressure 374
Edema 375Pressure-Volume Curve of the
Interstitial Fluid Spaces 375Positive Interstitial Fluid Pressure
as the Physical Basis forEdema 376
The Concept of a "Safety Factor"Before Edema Develops 376
Edema Resulting from AbnormalCapillary Dynamics 377
Edema Caused by Kidney Reten-tion of Fluid 379
The Presence and Importance of Gel inthe Interstitial Spaces 379
Pulmonary Interstitial Fluid Dynamics 380Pulmonary Edema 380
CHAPTER 32
THE SPECIAL FLUID SYSTEMS OFTHE BODY-CEREBROSPINAL,OCULAR, PLEURAL, PERICARDIAL,PERITONEAL, AND SYNOVIAL 383The Cerebrospinal Fluid System 383
Cushioning Function of the• Cerebrospinal Fluid 383Formation, Flow, and Absorption
of Cerebrospinal Fluid 384Cerebrospinal Fluid Pressure 385Obstruction to the Flow of
Cerebrospinal Fluid 385The Blood-Cerebrospinal Fluid
and Blood-Brain Barriers 385The Intraocular Fluid 386
Formation of Aqueous Humor bythe Ciliary Body 386
XVIII CONTENTS
Outflow of Aqueous Humor fromthe Eye 387
Intraocular Pressure 388Fluid Circulation in the Potential
Spaces of the Body - 389Fluid Exchange Between the
Capillaries and the PotentialSpaces 389
The Pleural Cavity 389The Pericardial Cavity 390The Peritoneal Cavity 390The Synovial Cavities 390
CHAPTER 33
PARTITION OF THE BODY FLUIDS:OSMOTIC EQUILIBRIA BETWEENEXTRACELLULAR AND INTRA-CELLULAR FLUIDS. 391
Total Body Water 391Intake Versus Output of Water 392
Body Fluid Compartments 392The Intracellular Compartment 392The Extracellular Fluid
Compartment 392Blood Volume 393
Measurement of Body Fluid Volumes 393The Dilution Principle for
Measuring Fluid Volumes 393Determination of Blood Volume 394Measurement of the Extracellular
Fluid Volume 395Measurement of Total Body
Water... 395Calculation of Interstitial Fluid
Volume.. , 396Constituents of Extracellular and
Intracellular Fluids 396Osmotic Equilibria and Fluid Shifts
Between the Extracellular and Intra-cellular Fluids 397.
Basic Principles of Osmosis andOsmotic Pressure 397
Osmolality of the Body Fluids 398Maintenance of Osmotic Equilib-
rium Between Extracellular andIntracellular Fluids 399
Changes in the Volumes and Osmolal-ities of the Extracellular and Intra-cellular Fluid Compartments inAbnormal States 400
Calculation of Fluid Shifts Be-tween the Extracellular andIntracellular Fluid Compart-ments 400
Effect of Adding Water to theExtracellular Fluid 400
Effect of Dehydration 401
Effect of Adding Saline Solutionto the Extracellular Fluid 401
Effect of Infusing HypertonicGlucose, Mannitol, or SucroseSolutions 401
Glucose and Other SolutionsAdministered for NutritivePurposes 402
CHAPTER 34
FORMATION OF URINE BY THEKIDNEY: GLOMERULAR FILTRATION,TUBULAR FUNCTION, AND PLASMACLEARANCE .; '. 403
Physiologic Anatomy of theKidney 403
Basic Theory of Nephron. Function 404
Renal Blood Flow and Pressures 405Blood Flow Through the Kidneys... 405Pressures in the Renal
Circulation 405"Intrarenal Pressure" and Renal
Interstitial Fluid Pressure 406Function of the Peritubular
Capillaries 406Glomerular Filtration and the
Glomerular Filtrate 406The Glomerular Filtration Rate 408Dynamics of Glomerular
Filtration 408Factors That Affect the Glomeru-
lar Filtration Rate 409Reabsorption and Secretion in the
Tubules 410Basic Mechanisms of Absorption
and Secretion in the Tubules 410Absorptive Capabilities of Dif-
ferent Tubule Segments 412Reabsorption and Secretion of
Individual Substances in Dif-ferent Segments of the Tubules ... 413
Concentrations of Different Sub-stances at Different Points inthe Tubules 416
The Concept of "Plasma Clearance" 417Inulin Clearance as a Measure of
Glomerular Filtration Rate 418Para-Aminohippuric Acid (PAH)
Clearance as a Measure ofPlasma Flow Through theKidneys 418
Calculating the Filtration Fractionfrom Plasma Clearances 418
Effect of "Tubular Load" and "TubularTransport Maximum" on UrineConstituents 418
CONTENTS XIX
CHAPTER 35
RENAL MECHANISMS FOR CONCEN-TRATING AND DILUTING THEURINE AND FOR UREAt SODIUM,POTASSIUM, AND FLUID VOLUMEEXCRETION 420Diluting Mechanism of the Kidney —The
Mechanism for Excreting Excess Water... 420Concentrating Mechanism of the
Kidney; Excretion of Excess Solutes —The Counter-Current Mechanism 421
Osmolar Clearance; Free WaterClearance 424
Urea Excretion 424Sodium Excretion 425Potassium Excretion 426Fluid Volume Excretion 427
1. Effect of Tubular OsmolarClearance on Rate of FluidVolume Excretion 428
2. Effect of Plasma.ColloidOsmotic Pressure on Rate ofFluid Volume Excretion 428
3. Effect of Sympathetic Stimula-tion on Rate of Fluid VolumeExcretion...; 429
4. Effect of Arterial Pressure onRate of Fluid Volume Excretion... 429
5. Effect of Antidiuretic Hormoneon Rate of Fluid VolumeExcretion 429
Summary of the Control of FluidVolume Excretion 430
Autoregulation of Glomerular Filtra-tion Rate 430
Autoregulation of Renal Blood Flow 432Role of the Renin-Angiotensin System
and of the Efferent VasoconstrictorMechanism in Conserving Water andSalt but Eliminating Urea DuringArterial Hypotension 433
CHAPTER 36
REGULATION OF BLOOD VOLUME,EXTRACELLULAR FLUID VOLUME,AND EXTRACELLULAR FLUIDCOMPOSITION BY THE KIDNEYSAND BY THE THIRST MECHANISM 435Control of Blood Volume 435
Basic Mechanism for BloodVolume Control 435
Role of the Volume Receptors inBlood Volume Control 436
Other Factors That Help toControl Blood Volume 437
Control of Extracellular Fluid Volume 438
Control of Extracellular Fluid SodiumConcentration and ExtracellularFluid Osmolality 438
The Osmo-Sodium Receptor-Antidiuretic Hormone Feed-back Control System 439
Thirst, and Its Role in ControllingSodium Concentration andOsmolality 441
Combined Roles of the Antidiu-retic and Thirst Mechanism forControl of Extracellular FluidSodium Concentration andOsmolality 442
Effect of Aldosterone on SodiumConcentration 443
Control of Sodium Intake —Appetite and Craving for Salt 443
Control of Extracellular Potassium Con-centration — Role of Aldosterone 444
Other Factors That Affect Potas-sium Ion Concentration 445
Control of the Extracellular Concen-trations of Other Ions 445
CHAPTER 37
REGULATION OF ACID-BASEBALANCE 448
Defense Against Changes in Hy-drogen Ion Concentration 448
Function of Acid-Base Buffers 449The Bicarbonate Buffer System ...:.. 449The Buffer Sys.tems of the Body
Fluids 450The Isohydric Principle 451
Respiratory Regulation of Acid-BaseBalance 452
Renal Regulation of Hydrogen IonConcentration 453
Tubular Secretion of HydrogenIons 453
Renal Correction of Alkalosis —Decrease in Bicarbonate Ionsin the Extracellular Fluid 455
Renal Correction of Acidosis —Increase in Bicarbonate Ions inthe Extracellular Fluid 455
Combination of the ExcessHydrogen Ions with TubularBuffers and Their Transportinto the Urine 456
Rapidity of Acid-Base Regulationby the Kidneys 457
Renal Regulation of PlasmaChloride Concentration —TheChloride to Bicarbonate Ratio 457
XX CONTENTS
Clinical Abnormalities of Acid-BaseBalance 458
Respiratory Acidosis andAlkalosis ."i '458
Metabolic Acidosis and Alkalosis... 458Effects of Acidosis and Alkalosis
on the Body 459Respiratory Compensation of
Metabolic Acidosis or Alkalosis... 459Renal Compensation of Respira-
tory Acidosis or Alkalosis 460Physiology of Treatment in
Acidosis or Alkalosis 460Clinical Measurements and
Analysis of Acid-BaseAbnormalities 460
CHAPTER 38
RENAL DISEASE, DIURESIS, ANDMICTURITION 463Renal Disease 463
Acute Renal Failure 463
Chronic Renal Failure —Decreasein Number of FunctionalNephrons 464
Hypertensive Kidney Disease 468The Nephrotic Syndrome —
Increased GlomerularPermeability 469
Specific Tubular Disorders 469Renal Function Tests 470Diuretics and Mechanisms of Their
Actions 470Micturition 471
Physiologic Anatomy of theBladder and Its NervousConnections ...'. 471
Transport of Urine Through theUreters 472
Tone of the Bladder Wall, and theCystometrogram DuringBladder Filling 472
The Micturition Reflex 473Abnormalities of Micturition 473
PART VIIRESPIRATION
CHAPTER 39
PULMONARY VENTILATION 476Mechanics of Pulmonary Ventilation 476
Basic Mechanisms of Lung Ex-pansion and Contraction' 476
Respiratory Pressures 477Expansibility of the Lungs and
Thorax: "Compliance" 478The "Work" of Breathing 479
The Pulmonary Volumes andCapacities 480
Recording Changes in PulmonaryVolume — Spirometry 480
The Pulmonary "Volumes" 480The Pulmonary "Capacities" 480Significance of the Pulmonary
-•Volumes and Capacities 481Abbreviations and Symbols Used
in Pulmonary Function Studies... 482Determination of Functional
Residual Capacity —The HeliumDilution Method 482
The Minute Respiratory Volume-Respiratory Rate and Tidal Volume 483
Forced Expiratory Vital Capacityand Forced ExpiratoryVolume '.:..' 484
Ventilation of the Alveoli 484The Dead Space 484Rate of Alveolar Ventilation 485
Functions of the Respiratory Passage-ways.. ' 486
Functions of the Nose 486The Cough Reflex '. 487The Sneeze Reflex 487Action of the Cilia to Clear
Respiratory Passageways 487Vocalization 487
Artificial Respiration 488Mouth-to-Mouth Breathing 488Mechanical Methods of Artificial
Respiration 488
CHAPTER 40
PHYSICAL PRINCIPLES OF GASEOUSEXCHANGE; DIFFUSION OF OXYGENAND CARBON DIOXIDE THROUGHTHE RESPIRATORY MEMBRANE
Physics of Diffusion and Gas Pressures ....The Molecular Basis of Gaseous
Diffusion
491491
491
CONTENTS XXI
Gas Pressures in a Mixture ofGases - Partial Pressures "ofIndividual Gases 492
Partial Pressure'of Gases in Waterand Tissues ' 492
The Vapor Pressure of Water 492Diffusion of Gases Through
Liquids —The Pressure Gradientfor Diffusion 493
Diffusion of Gases ThroughTissues 493
Composition of Alveolar Air1— ItsRelation to Atmospheric Air ^ 493
Rate at Which Alveolar Air IsRenewed by Atmospheric Air 494
Oxygen Concentration and PartialPressure in the Alveoli 495
CO 2 Concentration and PartialPressure in the Alveoli 495
Expired Air ..„. 496Diffusion of Gases Through the
Respiratory Membrane ...r 497Factors That Affect Rate of Gas
Diffusion Through,the Respira-tory, Membrane 498
Diffusing Capacity of the Respira-tory Membrane./ 499
Effect of the Ventilation-Perfusion Ratioon Alveolar Gas Concentration 500
CHAPTER 41
TRANSPORT OF OXYGEN ANDCARBON DIOXIDE IN THE BLOODAND BODY FLUIDS 504Pressures of Oxygen and Carbon
Dioxide in the Lungs, Blood, andTissues '. \ 504
Uptake of Oxygen by thePulmonary Blood 504
Transport of Oxygen in theArterial Blood : 505
Diffusion of Oxygen from theCapillaries to the InterstitialFluid 505
Diffusion of Oxygen from theCapillaries to the Cells 506
Diffusion of Carbon Dioxidefrom the Cells to the TissueCapillaries 506
Removal of Carbon Dioxide fromthe Pulmonary Blood 507
Transport of Oxygen in the Blood 507The Reversible Combination of
Oxygen with Hemoglobin 507The Oxygen Buffer Function of
Hemoglobin 509
Factors That Cause the Hemo-globin Dissociation Curve toShift 509
Total Rate of Oxygen Transport tfrom the Lungs to the Tissues 510
Metabolic Use of Oxygen by theCells 510
Transport of Oxygen in theDissolved State 511
Combination of Hemoglobinwith Carbon Monoxide 511
Transport of Carbon Dioxide in theBlood 512
Chemical Forms in Which CarbonDioxide Is Transported 512
The Carbon Dioxide DissociationCurve ., 513
Effect of the Oxygen-HemoglobinReaction on Carbon DioxideTransport-The Haldane Effect... 513
Change in Blood Acidity DuringCarbon Dioxide Transport 514
The Respiratory Exchange Ratio 514
CHAPTER 42
REGULATION OF RESPIRATION 516The Respiratory Center 516Chemical Control of Respiration 518
Direct Chemical Control ofRespiratory Center Activity byCarbon Dioxide and HydrogenIons 518
The Peripheral Chemoreceptor Systemfor Control of Respiratory Activity —Role of Oxygen in Respiratory Control... 520
Composite Effects of Pco2, pH,and Po2 on Respiratory Activity... 523
Regulation of Respiration DuringExercise 523
Other Factors That Affect Respiration 525Abnormalities of Respiratory Control 526
Respiratory Center Depression 526Periodic Breathing 526
CHAPTER 43
RESPIRATORY INSUFFICIENCY 529Additional Methods for Studying
Respiratory Abnormalities 529Study of Blood Gases and pH 530Physiologic Types of Respiratory
Insufficiency 531Abnormalities That Cause
Alveolar Hypoventilation 531Diseases That Decrease Lung
Diffusing Capacity 531
•v
XXII CONTENTS
Abnormalities of Oxygen Trans-port from the Lungs to theTissues 531
Physiologic Peculiarities of SpecificPulmonary Abnormalities 532
Chronic Emphysema 532Pneumonia 533Atelectasis 533Bronchial Asthma 534Tuberculosis 534
Hypoxia 535
Cyanosis 535Dyspnea 536
Hypercapnia 536Effects of Hypercapnia on the
Body 536Oxygen Therapy in the Different Types
of Hypoxia 536Danger of Hypercapnia During
Oxygen Therapy 538Absorption of Entrapped Air 538
PART VIMAVIATION, SPACE, AND DEEP SEA DIVING PHYSIOLOGY
CHAPTER 44
AVIATION, HIGH ALTITUDE, ANDSPACE PHYSIOLOGY..... 542Effects of Low Oxygen Pressure on the _
Body ..... 542Alveolar PO2 at Different
Elevations 7 542•/ Effect of Breathing Pure Oxygen <
on the Alveolar PO2 at DifferentAltitudes 543
The "Ceiling" When BreathingAir and When Breathing Oxygenin an Unpressurized Airplane 543
Effects of Hypoxia..; <-. 543Acclimatization to Low PO2 544Natural Acclimatization of
Natives Living at HighAltitudes 545
Work Capacity at High Altitudes:The Effect of Acclimatization 546
Chronic Mountain Sickness 546Effects of Acceleratory Forces on the
Body in Aviation and SpacePhysiology 546
Centrifugal Acceleratory Forces 547-
Effects of Linear AcceleratoryForces on the Body 548
Perceptions of Equilibrium andTurning in Blind Flying 549
Problems of Temperature in Aviationand Space Physiology 549
Radiation at High Altitudes and inSpace 550
"Artificial Climate" in the SealedSpacecraft 550
Weightlessness in Space 550
CHAPTER 45
PHYSIOLOGY OF DEEP SEA DIVINGAND OTHER HIGH-PRESSUREOPERATIONS 552Effect of High Partial Pressures of Gases
on the Body 552Decompression of the Diver After
Exposure to High Pressures 554Some Physical Problems of Diving 556Scuba Diving (Self-Contained Under-
water Breathing Apparatus) 557Special Physiological Problems in
Submarines 558
PART IXTHE NERVOUS SYSTEM
CHAPTER 46
ORGANIZATION OF THE NERVOUSSYSTEM; BASIC FUNCTIONS OF'SYNAPSES.......... 560
General Design of the Nervous System..... 560The Sensory Division—Sensory
Receptors 560The Motor Division —The
Effectors 560
I Processing of Information 561-Storage of Information—Memory ... 562
The Three Major Levels of NervousSystem Function 562
The Spinal Cord Level 562The Lower Brain Level 563The Higher Brain or Cortical
Level 563Comparison of the Nervous System with
an Electronic Computer 564
CONTENTS XXIII
Function of Central Nervous SystemSynapses 564
Physiologic Anatomy of theSynapse ." 565
Chemical and PhysiologicalNatures of the TransmitterSubstances 566
Electrical Events During NeuronalExcitation 568
Electrical Events in NeuronalInhibition 570
Summation of Postsynaptic :
Potentials 571Special Functions of Dendrites
in Exciting Neurons 572Relation of State of Excitation of
the Neuron to the Rate of Firing... 573Some Special Characteristics of Synaptic
Transmission 575
CHAPTER 47
^NEURONAL MECHANISMS ANDCIRCUITS FOR PROCESSINGINFORMATION ,.'..: 577
Information, Signals, andImpulses : ..... 577
Transmission of Signals in Nerve Tracts... 577Signal Strength .-..: 577Spatial Orientation of Signals in
Fiber Tracts 578Transmission and Processing of Signals
in Neuronal Pools -. c 579Transmission of Spatial Patterns
Through Successive NeuronalPools :.....' 581
Prolongation of a Signal by aNeuronal Pool—"After-Discharge" .'..! 583
Continuous Signal Output fromNeuronal Pools ,__.'. .' 585
Rhythmic Signal Output 585Instability and Stability of Neuronal
Circuits 586Inhibitory Circuits as a
Mechanism for StabilizingNervous System Function..... ~ 586
Synaptic Fatigue as a Means forStabilizing the Nervous System(Decremental Conduction) 586
CHAPTER 48
SENSORY RECEPTORS AND THEIRBASIC MECHANISMS OF ACTION 588Types of Sensory Receptors and the
Sensory Stimuli They Detect 588Differential Sensitivity "of
Receptors 588
Transduction of Sensory Stimuli intoNerve Impulses 590
Local Currents at Nerve Endings —Receptor Potentials andGenerator Potentials '. 590
Adaptation of Receptors 592Psychic Interpretation of Stimulus
Strength 593Judgment of Stimulus Strength 594
Physiological Classification of Nerve. Fibers 595
CHAPTER 49
SOMATIC SENSATIONS: I. THEMECHANORECEPTIVE SENSATIONS 597
Classification of Somatic Senses 597Detection and Transmission of Tactile
Sensations 597.- Detection of Vibration 599
The Subconscious "MuscleSense" ; 599
Tickling and Itch.... 599The Dual System for Transmission of
Mechanoreceptive Somatic SensorySignals into the Central NervousSystem 599
Function of the Spinal Cord Neuronsin Transmitting Sensory Signals 601
Transmission in the Dorsal-LemniscalSystem , '. 601
Anatomy of the Dorsal-LemniscalSystem 601
The Somesthetic Cortex 603Somatic Association Areas 605Characteristics of Transmission
in the Dorsal-Lemniscal System... 605The Position Sense 607
Transmission in the AnterolateralSpinothalamic System 608
Anatomy of the AnterolateralSpinothalamic Pathway 608
Some Special Aspects of SensoryFunction < 609
CHAPTER 50
SOMATIC SENSATIONS: II. PAIN,VISCERAL PAIN, HEADACHE, ANDTHERMAL SENSATIONS^ 611
Qualities of Pain 611Methods for Measuring the .-
Perception of Pain. 611The Pain Receptors and Their
Stimulation 612
XXIV CONTENTS
Rate of Tissue Damage as theCause of Pain 613
Transmission of Pain Signals into theCentral Nervous System 614
The Reaction to Pain and Its ControlWithin the Nervous System 615
A Pain Control ("Analgesic"). System in the Brain and SpinalCord 615
Referred Pain 617Referred Pain Caused by Reflex
Muscular Spasm 617Visceral Pain 617
Causes of True Visceral Pain 617"Parietal" Pain Caused by
Visceral Damage 618Localization of Visceral Pain—The
"Visceral" and the "Parietal"Transmission Pathways 618
Visceral Pain from Various•= Organs 619Some Clinical Abnormalities of Pain
and Other Sensations :.. 620Hyperalgesia ., 620The Thalamic Syndrome :..-....: 620Herpes Zoster... 621Tic Douloureux 621The Brown-Sequard Syndrome 621
Headache 621Headache of Intracranial Origin 621Extracranial Types of Headache 622
Thermal Sensations 623Thermal Receptors and Their
Excitation '. 623Transmission of Thermal Signals
in the Nervous System 624
CHAPTER 51 -
MOTOR FUNCTIONS OF THE SPINALCORD AND THE CORD REFLEXES....... 626
Organization of the Spinal Cordfor Motor Functions , 626
Role of the Muscle Spindle in MotorControl 628
Receptor Function of the MuscleSpindle 628
The Stretch Reflex (Also CalledMuscle Spindle Reflex orMyotatic Reflex)....: 629
Role of the Muscle Spindle in VoluntaryMotor Activity , 631
Clinical Applications of theStretch Reflex 632
The Tendon Reflex 633
The Flexor Reflex (the WithdrawalReflexes) 634
The Crossed Extensor Reflex 635Reciprocal Inhibition and Reciprocal
Innervation..-, 636Fatigue of Reflexes; Rebound 636The Reflexes of Posture and Locomotion... 636
The Postural and LocomotiveReflexes of the Cord 636
The Scratch Reflex 637The Spinal Cord Reflexes That Cause
Muscle Spasm 638The Autonomic Reflexes in the Spinal
Cord 638Spinal Cord Transection and Spinal
Shock 638
CHAPTER 52
MOTOR FUNCTIONS OF THE BRAINSTEM AND BASAL GANGLIA-RETICULAR FORMATION, VESTIBU-LAR APPARATUS, EQUILIBRIUM,AND BRAIN STEM REFLEXES 640The Reticular. Formation, and Support of
the Body Against Gravity 640Excitatory Function of the
Reticular Formation 641Support of the Body Against- Gravity 641
Vestibular Sensations and the Mainte-nance of Equilibrium 641
The Vestibular Apparatus 641Function of the Utricle and the
Saccule in the Maintenance ofStatic Equilibrium 643
The Semicircular Canals and Their De-tection of Angular Acceleration andAngular Velocity 644
Vestibular Postural Reflexes 645Vestibular Mechanism for
Stabilizing the Eyes and forNystagmus 645
Clinical Tests for Integrity ofVestibular Function 646
Other Factors Concerned withEquilibrium 646
Functions of the Reticular Formation andSpecific Brain Stem Nuclei in Con-trolling Subconscious, SterotypedMovements 647
Motor Functions of the Basal Ganglia 648Functions of the Different Basal
Ganglia 648Clinical Syndromes Resulting from
Damage to the Basal Ganglia 649
•v
CONTENTS xxv
CHAPTER 53 ; '
CORTICAL AND CEREBELLAR CON-TROL OF MOTOR FUNCTIONS.. 652Physiologic Anatomy of the Motor Areas
of the Cortex and Their Pathways tothe Cord 652
The Primary Motor Cortex of the .Human Being 654
Complex Movements Elicited byStimulating the CortexAnterior to the Motor Cortex—The Concept of a MotorAssociation Area 656
Effects of Lesions in the PrimaryMotor and Motor AssociationCortex 656
The Supplemental Motor Area 657Stimulation of the Spinal Motoneurons
by Motor Signals from the Brain...-. 657The Cerebellum and Its Motor Functions... 658
The Input System to theCerebellum .' ^ 659
Output Signals from the ..-Cerebellum : r 660
The Neuronal Circuit of theCerebellum.. 661
Function of the Cerebellum inControlling Movements 662
Function of the Cerebellum withthe Spinal Cord and LowerBrain Stem to Control Posturaland Equilibrium Movements 662
Clinical Abnormalities of theCerebellum 666
Sensory Feedback Control of MotorFunctions... 667
The Sensory Engram for MotorControl 668
Establishment of Rapid MotorPatterns 668
Initiation of Voluntary Motor 'Activity ; 669
CHAPTER 54
ACTIVATION OF THE BRAIN-THERETICULAR ACTIVATING SYSTEM;THE GENERALIZED THALAMOCOR-TICAL SYSTEM; BRAIN WAVES;EPILEPSY; WAKEFULNESS ANDSLEEP '. 671
Function of the Reticular Acti-vating System in Wakefulness..... 671
The Arousal Reaction—Activationof the Reticular ActivatingSystem by Sensory Signals 672
Stimulation of the ReticularActivating System by theCerebral Cortex 673
The Generalized.ThalamocorticalSystem 673
''Attention 674Possible "Searching" and "Pro-
gramming" Functions of theBrain's Activating System 675
Effect of Barbiturate Anesthesia onthe Reticular Activating System... 675
Brain Waves 675Origin of the Different Types of
Brain Waves 676Effect of Varying Degrees of
Cerebral Activity on the BasicRhythm of the Electroen-
y cephalogram 677Clinical Use of the Electroen-
cephalogram 677Epilepsy 677
Grand Mai Epilepsy 678Petit Mai Epilepsy 678Focal Epilepsy 679
Sleep and Wakefulness 679Slow Wave Sleep 679
' Paradoxical Sleep (Rem Sleep) 680Basic Theories of Sleep and
Wakefulness 680Physiological Effects of Sleep 682
CHAPTER 55
THE CEREBRAL CORTEX ANDINTELLECTUAL FUNCTIONS OFTHE BRAIN 684
Physiologic Anatomy of theCerebral Cortex 684
Functions of Certain Specific CorticalAreas.. .". 685
Specific Functions of the PrimarySensory Areas 685
The Sensory Association Areas 686Interpretative Function of the
Posterior Superior TemporalLobe —The General InterpretativeArea (or Wernicke's Area) 687
The Prefrontal Areas 689Thoughts, Consciousness, and Memory 690
Memory and Types of Memory L 690Physiological Basis of Memory 691
Analytical Operations of the Brain 694Function of the Brain in Communication... 695Function of the Corpus Callosum and
Anterior Commissure to TransferThoughts, Memories, and Other Infor-mation to the Opposite Hemisphere 697
•v
XXVI CONTENTS
CHAPTER 56
BEHAVIORAL FUNCTIONS OF THEBRAIN: THE LIMBIC SYSTEM, ROLEOF THE HYPOTHALAMUS, ANDCONTROL OF VEGETATIVE FUNC-TIONSOF THE BODY 699Functional Anatomy of the Limbic
System; Its Relation to theHypothalamus 699
The Hypothalamus, the Major OutputPathway of the Limbic System 700
Vegetative Control Functions ofthe Hypothalamus 701
Behavioral Functions of theHypothalamus and AssociatedLimbic Structures 703
Specific Functions of Other Parts of theLimbic System .- 705
Functions of the Amygdala 705Functions of the Hippocampus 705Function of the Limbic Cortex 706
Function of Specific Chemical Trans-mitter Systems for Behavior Control ...... 707
Psychosomatic Effects of the BehavioralSystem : 707
CHAPTER 57
THE AUTONOMIC NERVOUS SYSTEM;THE ADRENAL MEDULLA .... 710General Organization of the AutonomicNervous System .* 710
Physiologic Anatomy of theSympathetic Nervous System 710
Physiologic Anatomy of the Para-sympathetic Nervous System 711
Basic Characteristics of Sympatheticand Parasympathetic Function 712
Cholinergic and Adrenergic Fibers— Secretion of Acetylcholine L
or Norepinephrine by the Post-ganglionic Neurons 712
Receptor Substances of theEffector Organs ...-. 713
Excitatory and Inhibitory Actionsof Sympathetic and Parasympa-thetic Stimulation 714
Effects of Sympathetic and Para-sympathetic Stimulation onSpecific Organs 714
Function of the AdrenalMedullae 716
Relationship of Stimulus Rate toDegree of Sympathetic and Para-sympathetic Effect 717
Sympathetic and Parasympathetic"Tone" , 717
Denervation Supersensitivity ofSympathetic and Parasympa-thetic Organs Following
' c , Denervation 718The Autonomic Reflexes 718
Mass Discharge of the Sympa-thetic System Versus DiscreteCharacteristics of Para-sympathetic Reflexes 719
"Alarm" or "Stress" Function ofthe Sympathetic NervousSystem 719
° Medullary, Pontine, andMesencephalic Control of theAutonomic Nervous System 719
Pharmacology of the AutonomicNervous System 720
Drugs That Act on AdrenergicEffector Organs—The Sympa-thomimetic Drugs 720
Drugs That Act on CholinergicEffector Organs 720
,r' Drugs That Stimulate the Post-ganglionic Neurons—"NicotinicDrugs" 721
PART XTHE SPECIAL SENSES
CHAPTER 58
THE EYE: I. OPTICS OF VISION 724Physical Principles of Optics 724
Refraction of Light 724Application of Refractive
Principles to Lenses 724Focal Length of the Convex Lens 725Formation of an Image by a
Convex Lens 726
Measurement of the RefractivePower of a Lens —The Diopter 727
The Optics of the Eye 727The Eye as a Camera 727The Mechanism of Accommoda-
tion 728The Pupillary Aperture 729"Normal" Aberrations of Vision 730Errors of Refraction 731
CONTENTS XXVII
Size of the Image* on the Retinaand Visual Acuity 732
Determination of Distance of anObject from the Eye—DepthPerception 733
Optical Instruments 734The Ophthalmoscope 734The Retinoscope 734
CHAPTER 59
THE EYE: II. RECEPTOR FUNCTIONSOF THE RETINA 736Anatomy and Function of the Structural
Elements of the Retina.....^..:. 736Photochemistry of Visiond .-.r 739
The Rhodopsin-Retinal VisualCycle, and Excitation of theRods.... 739
Photochemistry of Color Visionby the Cones 741
Automatic Regulation of RetinalSensitivity—Dark and LightAdaptation .\ 742
Fusion of Flickering Lights bythe Retina : 743
Color Vision 744The Tri-Color Theory of Color
Perception....;.' 744Color Blindness "74"6
CHAPTER 60
THE EYE: III. NEUROPHYSIOLOGYOF VISION :,...- „ 748
The Visual Pathway 748Neural Function of the Retina 748
Neural Organization:of the Retina... 748Stimulation of the Rods and Cones
- The Receptor Potential 749Stimulation of the Bipolar and ,
Horizontal Cells ;.. 749Excitation of the Ganglion Cells 750Different Types of Signals Trans-
mitted by the Ganglion Cells,Through the Optic Nerve 750
Function of the Lateral Geniculate Body... 752Anatomical Organization of the
Lateral Geniculate Nuclei 752Function of the Primary Visual Cortex 753
Detection of Lines and Borders bythe Primary Visual Cortex 753
Perception of Luminosity 754Effect of Removing the Primary
Visual Cortex 754Transmission of Visual Informa-
tion into Other Regions of theCerebral Cortex ; 755
The Fields of Vision; Perimetry 755EyerMovements and Their Control 756
Conjugate Movement of the Eyes 757Fixation Movements of the Eyes 757Fusion of the Visual Images 759
Autonomic Control of Accommodationand Pupillary Aperture 760
Control of Accommodation 760, Control of the Pupillary Aperture... 760
CHAPTER 61
THE SENSE OF HEARING 763The Tympanic Membrane and the
Ossicular System 763Transmission of Sound from the
Tympanic Membrane to theCochlea 763
_ Transmission of Sound throughBone..... 764
The Cochlea 764Functional Anatomy of the
Cochlea 764Transmission of Sound Waves in
the Cochlea—The "TravelingWave" 766
Function of the Organ of Corti 767Determination of Pitch —The
"Place" Principle 768Determination of Loudhess 769
Central Auditory Mechanisms 770The Auditory Pathway 770Function of the Cerebral Cortex
in Hearing 771Discrimination of Direction from
Which Sound Emanates 772Centrifugal Conduction of Impulses
from the Central NervousSystem 773
Hearing Abnormalities 773Types of Deafness 773
CHAPTER 62
THE CHEMICAL SENSES-TASTEAND SMELL 775The Sense of Taste 775
The Primary Sensations of Taste 775The Taste Bud and Its Function ..... 776Transmission of Taste Signals into
the Central Nervous System 777Special Attributes of the Taste
Sense 778.The Sense of Smell v 779
The Olfactory Membrane 779Stimulation of the Olfactory Cells... 779Transmission of Smell Signals into D '•
the Central Nervous System 781
XXVIII CONTENTS
PART XITHE GASTROINTESTINAL TRACT
CHAPTER 63
MOVEMENT OF FOOD THROUGH THEALIMENTARY TRACT 784General Principles of Intestinal Motility... 784
Characteristics of the IntestinalWall 784
Innervation of the Gut —TheIntrinsic Nervous System 786
Hormonal Control of Gastro-intestinal Motility 787
Functional Types of Movements in theGastrointestinal Tract 787
The Mixing Movements 787The Propulsive Movements —
Peristalsis 787Ingestion of Food 788
Mastication (Chewing) 788Swallowing (Deglutition) 789Function of the Lower Esophageal
Sphincter (GastroesophagealSphincter) 791
Motor Functions of the Stomach 791Storage Function of the Stomach..... 791Mixing in the Stomach —The
Basic Electrical Rhythm of theStomach 792'
Emptying of the Stomach 793Movements of the Small Intestine 794
Mixing Contractions (Segmenta-tion Contractions) 795
Propulsive Movements 795Contraction and Emptying of the
Gallbladder— Cholecystokinin 796Function of the Ileocecal Valve 796
Movements of the Colon 797Defecation 798
Other Autonomic Reflexes AffectingBowel Activity 799
CHAPTER 64
SECRETORY FUNCTIONS OF THEALIMENTARY TRACT :.... 801General Principles of Gastrointestinal
Secretion 801Anatomical Types of Glands 801Basic Mechanisms of Stimulation
* of the Gastrointestinal Glands 801Basic Mechanism of Secretion by
Glandular Cells 802Lubricating and Protective
Properties of Mucus and ItsImportance in the Gastro-intestinal Tract 803
Secretion of Saliva 803Esophageal Secretion 805Gastric Secretion 805
Characteristics of the Gastric' Secretions 805
Regulation of Gastric Secretionby Nervous and HormonalMechanisms 807
Pancreatic Secretion 810Regulation of Pancreatic Secretion... 811
Secretion of Bile by the Liver 812Secretions of the Small Intestine 813
Secretion of Mucus by Brunner'sGlands and by Mucous Cells ofthe Intestinal Surface 813
Secretion of the Intestinal DigestiveJuices —The Crypts ofLieberkiihn 813
Regulation of Small IntestinalSecretion 814
Secretions of the Large Intestine 814
CHAPTER 65
DIGESTION AND ABSORPTION INTHEGASTROINTESTINAL TRACT 816Digestion of the Various Foods 816
, Digestion of Carbohydrates 816Digestion of Fats 817Digestion of Proteins 819
Basic Principles of GastrointestinalAbsorption 819
„ Anatomical Basis of Absorption 819Basic Mechanisms of Absorption ... 821
Absorption in the Small Intestine 821Absorption of Water 821Absorption of Ions 821Absorption of Nutrients •. 823
Absorption in the Large Intestine:^Formation of the Feces 825
CHAPTER 66
PHYSIOLOGY OF GASTRO-INTESTINAL DISORDERS 827Disorders of Swallowing and of the
Esophagus 827Disorders of the Stomach '. 827
Peptic Ulcer 828Disorders of the Small Intestine 830Disorders of the Large Intestine. 831
Constipation 831Diarrhea 831
CONTENTS XXIX
Paralysis of Defecation in SpinalCord Injuries 832
General Disorders of the GastrointestinalTract • 832
Vomiting 832
Nausea 833Gastrointestinal Obstruction :... 833Gases in the Gastrointestinal Tract
and Flatus 834
PART XIIMETABOLISM AND TEMPERATURE REGULATION
CHAPTER 67
METABOLISM OF CARBOHYDRATESAND FORMATION OF ADENOSINETRIPHOSPHATE 838
Release of Energy from Foods andthe Concept of "Free Energy" 838
Role of Adenosine Triphosphate "(ATP) in Metabolism 838
Central Role of Glucose in CarbohydrateMetabolism 839
Transport of Glucose Through the CellMembrane 840
Facilitation of Glucose Transportby Insulin 840
Phosphorylation of Glucose .\. 840Storage of Glycogen in Liver and Muscle... 840
Glycogenesis 840Glycogenolysis 841
Release of Energy from the GlucoseMolecule by the Glycolytic Pathway..... 841
Glycolysis and the Formation of, Pyruvic Acid 841Conversion of Pyruvic Acid to
Acetyl Coenzyme A 842The Citric Acid Cycle 842Formation of ATP by Oxidative
Phosphorylation 844Summary of ATP Formation During
the Breakdown of Glucose 845Control of Glycolysis and Oxida-
tion by Adenosine Diphosphate(ADP) 845
Anaerobic Release of Energy —"Anaerobic Glycolysis" 845
Release of Energy from Glucose by thePhosphogluconate Pathway 846
Glucose Conversion to Glycogenor Fat •* 847
Formation of Carbohydrates from Pro-teins and Fats—"Gluconeogenesis".. . 847
Blood Glucose 847
CHAPTER 68 ;
LIPID METABOLISM 849Transport of Lipids in the Blood 849
Transport from the GastrointestinalTract - The Chylomicrons 849
Transport of Fatty Acids in Combinationwith Albumin-Free Fatty Acid 850
The Lipoproteins 850The FatDeposits 851
Adipose Tissue 851The Liver Lipids 851
Use of Triglycerides for Energy, andFormation of Adenosine Triphosphate(ATP) 851
Formation of Acetoacetic Acid in({he Liver and Its Transport in
•the Blood 853Synthesis of Triglycerides from
Carbohydrates 853Synthesis of Triglycerides from
Proteins : 855Regulation of Energy Release from
Triglycerides 855Obesity 856
Phospholipids and Cholesterol 856Phospholipids 856Cholesterol 857
_v Structural Functions —Especiallyfor Membranes — of Phospho-lipids and Cholesterol 858
Atherosclerosis 858Experimental Production of
Atherosclerosis in Animals 858Atherosclerosis 858
Atherosclerosis in the HumanBeing 859
Summary of Factors CausingAtherosclerosis 859
CHAPTER 69
PROTEIN METABOLISM 861Basic Properties of Proteins 861
The Amino Acids 861Physical Characteristics of Proteins 861
Globular Proteins 861Fibrous Proteins 862Conjugated Proteins 863
Transport and Storage of Amino Acids 863The Blood Amino Acids 863Storage of Amino Acids as Proteins
in the Cells 863The Plasma Proteins 864Chemistry of Protein Synthesis 865
•v
XXX CONTENTS
Use of Proteins for Energy 865Obligatory Degradation of
Proteins 866Hormonal Regulation of Protein >
Metabolism 867
CHAPTER 70
THE LIVER AND BILIARY SYSTEM 869Physiologic Anatomy of the Liver... 869
Function of the Hepatic VascularSystem 870
Secretion of Bile and Functions of theBiliary Tree 870
Physiologic Anatomy of BiliarySecretion '. 870
The Bile Salts and Their Function... 871Excretion of Bilirubin in the Bile ... 872Secretion of Cholesterol; Gallstone
Formation 874Metabolic Functions of the Liver 8 74
Carbohydrate Metabolism 875Fat Metabolism 875'Protein Metabolism 875Miscellaneous Metabolic Func-
tions of the Liver 876
CHAPTER 71
ENERGETICS AND METABOLICRATE..... 877Importance of Adenosine Triphosphate
(ATP) in Metabolism 877Creatine Phosphate as a Storage .
Depot for Energy : 878Anaerobic Versus Aerobic Energy... 878Summary of Energy Utilization by
the Cells 879Control of Energy Release in the Cell 880The Metabolic Rate 881
Measurement of the MetabolicRate 881
Factors That Affect the MetabolicRate 882
The Basal Metabolic Rate 884
CHAPTER. 72
BODY TEMPERATURE, TEMPERATUREREGULATION, AND FEVER 886
The Insulator System of the Body... 886Flow of Blood to the Skin and Heat ;
Transfer from the Body Core 887Balance Between Heat Production and
Heat Loss .' 887Heat Loss 887
Sweating and Its Regulation bythe Autonomic Nervous System... 889
Regulation of Body Temperature—The"Hypothalamic Thermostat" 890
Neuronal Effector Mechanisms to'Increase or Decrease BodyTemperature , 891
The Concept of a "Set-Point" forTemperature Control 893
Behavioral Control of BodyTemperature 894
Local Skin Reflexes 894Abnormalities of Body Temperature
Regulation , 894Fever 894Exposure of the Body to Extreme
Cold 897
CHAPTER 73
DIETARY BALANCES, REGULATIONOF FEEDING; OBESITY ANDSTARVATION 899Dietary Balances 899
Energy Available in Foods 899Average Composition of the Diet 899Study of Energy Balances 900
Regulation of Food Intake 902Neural Centers for Regulation of
Food Intake 902Factors That Regulate Food Intake... 902
Obesity:- .?.' 904Treatment of Obesity 905
Inanition 905Starvation 905
CHAPTER 74
VITAMIN AND MINERALMETABOLISM 907Vitamins 907
Vitamin A.. 907thiamine (Vitamin Bi)..... 908Niacin 908Riboflavin (Vitamin B2) 909Vitamin B12 909Folic Acid (Pteroylglutamic Acid)... 910
V Pyridoxine (Vitamin B6) 9107 ^ Pantothenic Acid 910
Ascorbic Acid (Vitamin C) 910Vitamin D 911Vitamin E 911Vitamin K 912
Mineral Metabolism 912
CONTENTS XXXI
PART XIIIENDOCRINOLOGY AND REPRODUCTION
CHAPTER 75
INTRODUCTION TOENDOCRINOLOGY; AND THEPITUITARY HORMONES 916
Nature of a Hormone 916Mechanisms of Hormonal Action 917
Intracellular Hormonal MediatorsCyclic AMP 918
Action of Steroid Hormones onthe Genes to Cause ProteinSynthesis 918
Other Mechanisms of HormoneFunction ? 919
The Pituitary Gland and Its Relationshipto the Hypothalamus ' 919
Control of Pituitary Secretion bythe Hypothalamus 920
The Anterior Pituitary Gland and ItsRegulation by Hypothalamic ReleasingFactors 920
Cell Types of the AnteriorPituitary 920
The Hypothalamic-HypophysialPortal System 920
Physiological Functions of the AnteriorPituitary Hormones 921
Growth Hormone 921Abnormalities of Growth Hormone
Secretion 925The Posterior Pituitary Gland and Its
Relation to the Hypothalamus 927Physiological Functions of Anti-
diuretic Hormone (Vasopressin)... 927Oxytocic Hormone 929Chemical Nature of Antidiuretic
Hormone (Vasopressin) andOxytocin 929
CHAPTER 76
THE THYROID HORMONES : 931Formation and Secretion of the Thyroid >
Hormones 931Requirements of Iodine for
Formation of Thyroxine 931 ̂The Iodide Pump (Iodide
Trapping) 932Thyroglobulin and Chemistry of
Thyroxine and Triiodothyronine,Formation 932
Release of Thyroxine and Triiodo-thyronine from Thyroglobulin 933
Transport of Thyroxine and Tri-iodothyronine to the Tissues 933
Functions of the Thyroid Hormones inthe Tissues 934
General Increase in Metabolic v
Rate 934Effect of Thyroid Hormone on
Growth 935Effects of Thyroid Hormone on
Specific Bodily Mechanisms 935Regulation of Thyroid Hormone
Secretion .' 937Antithyroid Substances 939
Diseases of the Thyroid 939Hyperthyroidism 939Hypothyroidism 941
CHAPTER 77
THE ADRENOCORTICALHORMONES 944Functions of the Mineralocorticoids —
Aldosterone 944Renal Effects of Aldosterone 945Effects of Aldosterone on Sweat
Glands, Salivary Glands, andIntestinal Absorption 946
Cellular Mechanism ofAldosterone Action 946
• Regulation of AldosteroneSecretion 946
Functions of the Glucocorticoids 948Effect of Cortisol on Carbohydrate
Metabolism 948Effects of Cortisol on Protein
Metabolism 949Effects of. Cortisol on Fat
Metabolism ! 949Other Effects of Cortisol 950Regulation of Cortisol Secretion—
Adrenocorticotropic Hormone(ACTH) 951
Chemistry of Adrenocortical Secretion 953The Adrenal Androgens 954Abnormalities of Adrenocortical
Secretion 955Hypoadrenalism—Addison's
Disease 955Hyperadrenalism—Cushing's
Syndrome 955Primary Aldosteronism 956Adrenogenital Syndrome 957
XXXII CONTENTS
CHAPTER 78
INSULIN, GLUCAGON, ANDDIABETES MELLITUS.... :.'..'.. 959The Metabolic Effects of Insulin 959
Effect of Insulin on CarbohydrateMetabolism , 960
Effect of Insulin on FatMetabolism 962
Effect of Insulin on ProteinMetabolism and Growth 964
Control of Insulin Secretion 965Role of Insulin in "Switching"
Between Carbohydrate andLipid Metabolism 966
Glucagon and Its Functions 966Regulation of Glucagon Secretion... 967Glucagon-Like Effects of
Epinephrine 967Summary of Blood Glucose Regulation 968Diabetes Mellitus 968
Pathological Physiology ofDiabetes 969
Physiology of Diagnosis 969Treatment of Diabetes 970Diabetic Coma ~ 971
Hyperinsulinism 971
CHAPTER 79 „
PARATHYROID HORMONE, CALCI-TONIN, CALCIUM AND PHOSPHATEMETABOLISM, VITAMIN D, BONE,AND TEETH 973Calcium and Phosphate in the Extra-
cellular Fluid and Plasma —Functionof Vitamin D 973
Absorption and Excretion ofCalcium and Phosphate 973
Vitamin D and Its Role in Calciumand Phosphate Absorption 974
The Calcium in the Plasma andInterstitial Fluid 976
The Inorganic Phosphate in theExtracellular Fluids 976
Effects of Altered Calcium and -', <_.\ Phosphate Concentrations in
the Body Fluids 976Bone and Its Relationships to Extra-
cellular Calcium and Phosphates 977Precipitation and Absorption of
Calcium and Phosphate in Bone— Equilibrium with the Extra-cellular Fluids 978
Exchangeable Calcium 979Deposition and Absorption of
Bone —Remodeling of Bone 979
Parathyroid Hormone 981Effect of Parathyroid Hormone on
Calcium and Phosphate Concen-trations in the ExtracellularFluid :. 981
Control of Parathyroid Secretion byCalcium Ion Concentration 983
Calcitonin....^ 984Overall Control of Calcium Ion
Concentration 985Physiology of Parathyroid and Bone
Diseases 986Hypoparathyroidism 986Hyperparathyroidism 986Rickets 987Osteoporosis 988
Physiology of the Teeth 988Function of the Different Parts
of the Teeth 988Dentition 989Mineral Exchange in Teeth 990Dental Abnormalities 990
1 i
CHAPTER 80
REPRODUCTIVE AND HORMONALFUNCTIONS OF THE MALE; ANDTHE PINEAL GLAND 992Spermatogenesis : 992
The Steps of Spermatogenesis 992The Sperm 993Function of the Seminal Vesicles ... 994Function of the Prostate Gland 994Semen., 994Male Fertility 995
The Male Sexual Act 996Neuronal Stimulus for Performance
of the Male SexualAct 996Stages of the Male Sexual Act 996
Testosterone and Other Male SexHormones 997
Secretion, Metabolism, andChemistry of the Male SexHormone 997
Functions of Testosterone 998Basic Intracellular Mechanism of
Action of Testosterone 1000Control of Male Sexual Functions
by the Gonadotropic Hormones-FSH and LH 1000
Abnormalities of Male SexualFunction 1002
The Prostate-Gland and ItsAbnormalities 1002
Hypogonadism in the Male 1002Testicular Tumors and Hyper-
gonadism in the Male 1002
CONTENTS XXXIII
The Pineal Gland —Its Function inControlling Seasonal Fertility 1003
CHAPTER 81
PREPREGNANCY REPRODUCTIVEFUNCTIONS OF THE FEMALE, ANDTHE FEMALE HORMONES 1005Physiologic Anatomy of the Female
Sexual Organs 1005The Female Hormonal System 1005The Monthly Ovarian Cycle and
Function of the Gonadotropic c
Hormones 1006Follicular Growth —Function of
Follicle-Stimulating Hormone(FSH) 1006
The Corpus Luteum—the "Luteal"Phase of the Ovarian Cycle 1008
Summary 1009Functions of the Ovarian Hormones —
Estrogens and Progesterone 1009Chemistry of the Sex Hormones 1009Functions of the Estrogens —
Effects on the Primary andSecondary SexualCharacteristics ;...... 1011
Functions of Progesterone 1012The Endometrial Cycle and
Menstruation ..1013Regulation of the Female Monthly
Rhythm—Interplay Between theOvarian and Hypothalamic-PituitaryHormones 1014
Feedback Oscillation of theHypothalamic-Pituitary-Ovarian
* System 1015I Puberty and Menarche 1015< The Menopause 1016I Interrelationships of the Ovaries with -I OtherGlands ,>... 1016i Abnormalities of Secretion by the ' ^
Ovaries ,.'.... v....'. 1017The Female Sexual Act 1017Female Fertility '..*: 1018
CHAPTER 82
PREGNANCY AND LACTATION 1021Maturation of the Ovum 1021Fertilization of the Ovum 1021Transport and Implantation of the
Developing Ovum 1022Transport of the Ovum Through ,-
the Fallopian Tube \... 1022Implantation of the Ovum in the ^
Uterus 1022
Early Intrauterine Nutrition of theEmbryo 1023
Function of the Placenta 1023Developmental and Physiologic
Anatomy of the Placenta 1023Permeability of the Placental
Membrane 1024Storage Function of the Placenta 1026
Hormonal Factors in Pregnancy 1026Human Chorionic Gonadotropin
and Its Effect in CausingPersistence of the CorpusLuteum and in PreventingMenstruation 1026
Secretion of Estrogens by thePlacenta 1027
Secretion of Progesterone by thePlacenta 1027
Human Chorionic Somatomam-motropin 1027
Other Hormonal Factors inPregnancy 1028
Response of the Mother's Body toPregnancy 1028
Changes in the Maternal Circula-tory System During Pregnancy 1029
Weight Gain in the PregnantWoman..... 1029
Metabolism During Pregnancy 1029Respiration During Pregnancy 1030Function of the Maternal Urinary
System During Pregnancy 1030The Amniotic Fluid and Its
Formation 1030Abnormal Responses to Pregnancy 1030
Hyperemesis Gravidarum 1030i Preeclampsia and Eclampsia 1030
Parturition 1031Increased Uterine Irritability Near
Term 1031Onset of Labor—A Positive Feed-
back Theory for Its Initiation 1031Abdominal Muscle Contraction
During Labor 1032Mechanics of Parturition 1032Separation and Delivery of the .
Placenta J 1033Labor Pains 1033Involution of the Uterus 1033
Lactation 1033Development of the Breasts 1033Initiation of Lactation —Function
of Prolactin 1034The Ejection or "Let-Down"
Process in Milk Secretion—Function of Oxytocin 1035
Milk and the Metabolic Drain onthe Mother Caused by Lactation... 1035
XXXIV CONTENTS
CHAPTER 83
SPECIAL FEATURES OF FETAL ANDNEONATAL PHYSIOLOGY 1037Growth and Functional Developmentof the Fetus 1037
Development of the OrganSystems 1037
Function of the Nervous System 1038Function of the Gastrointestinal
Tract 1038Function of the Kidneys 1038Metabolism in the Fetus : 1038
Adjustments of the Infant toExtrauterine Life 1039
Onset of Breathing 1039
Circulatory Readjustments atBirth... 1040
Nutrition of the Newborn Infant 1041Special Functional Problems in the
Neonatal Infant T : 1041Special Problems of Prematurity 1044
Immature Development of thePremature Infant 1044
Instability of the Control Systemsin the Premature Infant 1044
Danger of Oxygen Therapy in thePremature Infant 1045
Growth and Development of the Child 1045Behavioral Growth 1045
INDEX 1047