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REVIEW OF MEDICAL PHYSIOLOGY - 20th Ed. (2001)Front MatterReadmeSection I. IntroductionSection II. Physiology of Nerve & Muscle CellsSection III. Functions of the Nervous SystemSection IV. Endocrinology, Metabolism, & Reproductive FunctionSection V. Gastrointestinal FunctionSection VI. CirculationSection VII. RespirationSection VIII. Formation & Excretion of UrineAppendixSelf-StudyAnswers

REVIEW OF MEDICAL PHYSIOLOGY - 20th Ed. (2001)Front MatterTitle Pagea LANGE medical bookReview of Medical Physiologytwentieth editionWilliam F. Ganong, MDJack and DeLoris Lange Professor of Physiology EmeritusUniversity of CaliforniaSan FranciscoLange Medical Books/McGraw-HillMedical Publishing DivisionNew York Chicago San Francisco Lisbon London Madrid Mexico City MilanNew Delhi San Juan Seoul Singapore Sydney TorontoCopyright PageReview of Medical Physiology, Twentieth EditionCopyright 2001 by The McGraw-Hill Companies, Inc. All rights reserved. Printed in the United Statesof America. Except as permitted under the United States Copyright Act of 1976, no part of this publicationmay be reproduced or distributed in any form or by any means, or stored in a data base or retrievalsystem, without the prior written permission of the publisher.Previous editions copyright 1999, 1997, 1995, 1993, 1991, by Appleton & Lange; copyright 1963through 1989 by Lange Medical Publications.1 2 3 4 5 6 7 8 9 0 DOW/DOW 0 9 8 7 6 5 4 3 2 1ISBN 0-8385-8282-6ISSN 0892-1253Notice: Medicine is an ever-changing science. As new research and clinical experience broaden ourknowledge, changes in treatment and drug therapy are required. The author and the publisher of thiswork have checked with sources believed to be reliable in their efforts to provide information that iscomplete and generally in accord with the standards accepted at the time of publication. However, in viewof the possibility of human error or changes in medical sciences, neither the author nor the publisher norany other party who has been involved in the preparation or publication of this work warrants that theinformation contained herein is in every respect accurate or complete, and they disclaim all responsibilityfor any errors or omissions or for the results obtained from use of the information contained in this work.Readers are encouraged to confirm the information contained herein with other sources. For exampleand in particular, readers are advised to check the product information sheet included in the package ofeach drug they plan to administer to be certain that the information contained in this work is accurate andthat changes have not been made in the recommended dose or in the contraindications foradministration. This recommendation is of particular importance in connection with new or infrequentlyused drugs.The book was set in Times Roman by Rainbow Graphics.

The editors were Janet Foltin, Isabel Nogueira, Jim Ransom, and Lester A. Sheinis.The production supervisor was Phil Galea.The production service was Rainbow Graphics.The cover designer was Mary McKeon.The art manager was Charissa Baker.The art coordinator was Becky Hainz-Baxter.The illustrators were Linda F. Harris, Shirley Bortoli, and Teshin Associates.The indexer was Katherine Pitcoff.R. R. Donnelley & Sons Company was printer and binder.This book is printed on acid-free paper.INTERNATIONAL EDITION ISBN 0-07-112064-5Copyright 2001. Exclusive rights by The McGraw-Hill Companies, Inc., for manufacture and export.This book cannot be re-exported from the country to which it is consigned by McGraw-Hill. TheInternational Edition is not available in North America.ContentsPreface ixSECTION I. INTRODUCTION 11. The General & Cellular Basis of Medical Physiology 1Introduction 1General Principles 1Functional Morphology of the Cell 8Structure & Function of DNA & RNA 17Transport Across Cell Membranes 27The Capillary Wall 35Intercellular Communication 35Homeostasis 46Aging 46Section I References: 47SECTION II. PHYSIOLOGY OF NERVE & MUSCLE CELLS 472. Excitable Tissue: Nerve 49Introduction 49Nerve Cells 49Excitation & Conduction 51Ionic Basis of Excitation & Conduction 56Properties of Mixed Nerves 58Nerve Fiber Types & Function 58Neurotrophins 58Glia 613. Excitable Tissue: Muscle 62Introduction 62Skeletal Muscle 62Morphology 62Electrical Phenomena & Ionic Fluxes 65Contractile Responses 65Energy Sources & Metabolism 70Properties of Muscles in the Intact Organism 72Cardiac Muscle 74Morphology 74

Electrical Properties 74Mechanical Properties 75Metabolism 77Pacemaker Tissue 78Smooth Muscle 78Morphology 78Visceral Smooth Muscle 78Multi-Unit Smooth Muscle 804. Synaptic & Junctional Transmission 81Introduction 81Synaptic Transmission 81Functional Anatomy 81Electrical Events in Postsynaptic Neurons 84Inhibition & Facilitation at Synapses 88Chemical Transmission of Synaptic Activity 90Principal Neurotransmitter Systems 93Synaptic Plasticity & Learning 110Neuromuscular Transmission 110Neuromuscular Junction 110Nerve Endings in Smooth & Cardiac Muscle 112Denervation Hypersensitivity 1135. Initiation of Impulses in Sense Organs 115Introduction 115Sense Organs & Receptors 115The Senses 115Electrical & Chemical Events in Receptors 117Ionic Basis of Excitation 118"Coding" of Sensory Information 119Section II References: 121SECTION III. FUNCTIONS OF THE NERVOUS SYSTEM 1236. Reflexes 123Introduction 123Monosynaptic Reflexes: The Stretch Reflex 123Polysynaptic Reflexes: The Withdrawal Reflex 129General Properties of Reflexes 1307. Cutaneous, Deep, & Visceral Sensation 132Introduction 132Pathways 132Touch 135Proprioception 136Temperature 136Pain 136Other Sensations 1428. Vision 144Introduction 144Anatomic Considerations 144The Image-Forming Mechanism 149The Photoreceptor Mechanism 152Responses in the Visual Pathways & Cortex 155Color Vision 159Other Aspects of Visual Function 161

Eye Movements 1639. Hearing & Equilibrium 166Introduction 166Anatomic Considerations 166Hair Cells 170Hearing 172Vestibular Function 17810. Smell & Taste 180Introduction 180Smell 180Taste 183Receptor Organs & Pathways 18311. Alert Behavior, Sleep, & the Electrical Activity of the Brain 187Introduction 187The Thalamus & the Cerebral Cortex 187The Reticular Formation & the Reticular Activating System 187Evoked Cortical Potentials 188The Electroencephalogram 189Physiologic Basis of the EEG, Consciousness, & Sleep 19112. Control of Posture & Movement 197Introduction 197General Principles 197Corticospinal & Corticobulbar System 198Anatomy & Function 198Posture-Regulating Systems 201Spinal Integration 203Medullary Components 204Midbrain Components 206Cortical Components 207Basal Ganglia 207Cerebellum 21113. The Autonomic Nervous System 217Introduction 217Anatomic Organization of Autonomic Outflow 217Chemical Transmission at Autonomic Junctions 219Responses of Effector Organs to Autonomic Nerve Impulses 22114. Central Regulation of Visceral Function 224Introduction 224Medulla Oblongata 224Hypothalamus 225Anatomic Considerations 225Hypothalamic Function 226Relation to Autonomic Function 226Relation to Sleep 227Relation to Cyclic Phenomena 227Hunger 228Thirst 232Control of Posterior Pituitary Secretion 233Control of Anterior Pituitary Secretion 239Temperature Regulation 24215. Neural Basis of Instinctual Behavior & Emotions 248

Introduction 248Anatomic Considerations 248Limbic Functions 249Sexual Behavior 249Fear & Rage 252Motivation & Addiction 253Brain Chemistry & Behavior 25416. "Higher Functions of the Nervous System": Conditioned Reflexes, Learning, & RelatedPhenomena 259Introduction 259Methods 259Learning & Memory 259Functions of the Neocortex 264Section III References: 264SECTION IV. ENDOCRINOLOGY, METABOLISM, & REPRODUCTIVE FUNCTION 26717. Energy Balance, Metabolism, & Nutrition 271Introduction 271Energy Metabolism 271Intermediary Metabolism 274Carbohydrate Metabolism 278Protein Metabolism 284Fat Metabolism 290Nutrition 30218. The Thyroid Gland 307Introduction 307Anatomic Considerations 307Formation & Secretion of Thyroid Hormones 308Transport & Metabolism of Thyroid Hormones 311Effects of Thyroid Hormones 313Regulation of Thyroid Secretion 316Clinical Correlates 31719. Endocrine Functions of the Pancreas & Regulation of Carbohydrate Metabolism 322Introduction 322Islet Cell Structure 322Structure, Biosynthesis, & Secretion of Insulin 323Fate of Secreted Insulin 324Effects of Insulin 324Mechanism of Action 327Consequences of Insulin Deficiency 328Insulin Excess 333Regulation of Insulin Secretion 334Glucagon 337Other Islet Cell Hormones 339Effects of Other Hormones & Exercise on Carbohydrate Metabolism 340Hypoglycemia & Diabetes Mellitus in Humans 34120. The Adrenal Medulla & Adrenal Cortex 344Introduction 344Adrenal Morphology 344Adrenal Medulla 346Structure & Function of Medullary Hormones 346Regulation of Adrenal Medullary Secretion 348

Adrenal Cortex 349Structure & Biosynthesis of Adrenocortical Hormones 349Transport, Metabolism, & Excretion of Adrenocortical Hormones 354Effects of Adrenal Androgens & Estrogens 356Physiologic Effects of Glucocorticoids 356Pharmacologic & Pathologic Effects of Glucocorticoids 358Regulation of Glucocorticoid Secretion 359Effects of Mineralocorticoids 362Regulation of Aldosterone Secretion 364Role of Mineralocorticoids in the Regulation of Salt Balance 366Summary of the Effects of Adrenocortical Hyper- & Hypofunction in Humans 36621. Hormonal Control of Calcium Metabolism & the Physiology of Bone 369Introduction 369Calcium & Phosphorus Metabolism 369Bone Physiology 370Vitamin D & the Hydroxycholecalciferols 375The Parathyroid Glands 377Calcitonin 380Effects of Other Hormones & Humoral Agents on Calcium Metabolism 38222. The Pituitary Gland 383Introduction 383Morphology 381Intermediate-Lobe Hormones 384Growth Hormone 386Physiology of Growth 392Pituitary Insufficiency 395Pituitary Hyperfunction in Humans 39623. The Gonads: Development & Function of the Reproductive System 398Introduction 398Sex Differentiation & Development 398Chromosomal Sex 398Embryology of the Human Reproductive System 400Aberrant Sexual Differentiation 401Puberty 405Precocious & Delayed Puberty 407Menopause 408Pituitary Gonadotropins & Prolactin 408The Male Reproductive System 410Structure 410Gametogenesis & Ejaculation 411Endocrine Function of the Testes 415Control of Testicular Function 418Abnormalities of Testicular Function 419The Female Reproductive System 419The Menstrual Cycle 419Ovarian Hormones 425Control of Ovarian Function 430Abnormalities of Ovarian Function 433Pregnancy 433Lactation 43624. Endocrine Functions of the Kidneys, Heart, & Pineal Gland 439Introduction 439The Renin-Angiotensin System 439Erythropoietin 444

Hormones of the Heart & Other Natriuretic Factors 445Pineal Gland 447Section IV References: 449SECTION V. GASTROINTESTINAL FUNCTION 45325. Digestion & Absorption 453Introduction 453Carbohydrates 453Proteins & Nucleic Acids 456Lipids 458Absorption of Water & Electrolytes 459Absorption of Vitamins & Minerals 46226. Regulation of Gastrointestinal Function 464Introduction 464General Considerations 464Gastrointestinal Hormones 466Mouth & Esophagus 472Stomach 475Exocrine Portion of the Pancreas 481Liver & Biliary System 483Small Intestine 489Colon 492Section V References: 496SECTION VI. CIRCULATION 49927. Circulating Body Fluids 499Introduction 499Blood 499Bone Marrow 499White Blood Cells 500Immunity 504Platelets 514Red Blood Cells 515Blood Types 519Plasma 522Hemostasis 524Lymph 52728. Origin of the Heartbeat & the Electrical Activity of the Heart 528Introduction 528Origin & Spread of Cardiac Excitation 528The Electrocardiogram 530Cardiac Arrhythmias 535Electrocardiographic Findings in Other Cardiac & Systemic Diseases 54129. The Heart as a Pump 545Introduction 545Mechanical Events of the Cardiac Cycle 545Cardiac Output 55030. Dynamics of Blood & Lymph Flow 556Introduction 556Anatomic Considerations 556

Biophysical Considerations 560Arterial & Arteriolar Circulation 565Capillary Circulation 568Lymphatic Circulation & Interstitial Fluid Volume 570Venous Circulation 57231. Cardiovascular Regulatory Mechanisms 574Introduction 574Local Regulatory Mechanisms 574Substances Secreted by the Endothelium 575Systemic Regulation by Hormones 577Systemic Regulation by the Nervous System 57932. Circulation Through Special Regions 588Introduction 588Cerebral Circulation 588Anatomic Considerations 588Cerebrospinal Fluid 589The Blood-Brain Barrier 591Cerebral Blood Flow 593Regulation of Cerebral Circulation 595Brain Metabolism & Oxygen Requirements 596Coronary Circulation 597Splanchnic Circulation 601Circulation of the Skin 602Placental & Fetal Circulation 60333. Cardiovascular Homeostasis in Health & Disease 607Introduction 607Compensations for Gravitational Effects 607Exercise 609Inflammation & Wound Healing 612Shock 613Hypertension 618Heart Failure 620Section VI References: 622SECTION VII. RESPIRATION 62534. Pulmonary Function 625Introduction 625Properties of Gases 625Anatomy of the Lungs 626Mechanics of Respiration 627Gas Exchange in the Lungs 637Pulmonary Circulation 639Other Functions of the Respiratory System 64235. Gas Transport Between the Lungs & the Tissues 644Introduction 644Oxygen Transport 644Carbon Dioxide Transport 64736. Regulation of Respiration 649Introduction 649Neural Control of Breathing 649Regulation of Respiratory Activity 650

Chemical Control of Breathing 651Nonchemical Influences on Respiration 65637. Respiratory Adjustments in Health & Disease 658Introduction 658Effects of Exercise 658Hypoxia 660Hypoxic Hypoxia 661Other Forms of Hypoxia 667Oxygen Treatment 668Hypercapnia & Hypocapnia 668Other Respiratory Abnormalities 669Effects of Increased Barometric Pressure 670Artificial Respiration 672Section VII References: 673SECTION VIII. FORMATION AND EXCRETION OF URINE 67538. Renal Function & Micturition 675Introduction 675Functional Anatomy 675Renal Circulation 679Glomerular Filtration 681Tubular Function 684Water Excretion 689Acidification of the Urine & Bicarbonate Excretion 694Regulation of Na & Cl - Excretion 697Regulation of K Excretion 699Diuretics 699Effects of Disordered Renal Function 700Filling of the Bladder 701Emptying of the Bladder 70139. Regulation of Extracellular Fluid Composition & Volume 704Introduction 704Defense of Tonicity 704Defense of Volume 704Defense of Specific Ionic Composition 705Defense of H Concentration 705Section VIII References: 713Appendix 714General References 714Normal Values & the Statistical Evaluation of Data 714Abbreviations & Symbols Commonly Used in Physiology 716Some Standard Respiratory Symbols 722Equivalents of Metric, United States, & English Measures 723Greek Alphabet 723Self-Study: Objectives, Essay Questions, & Multiple-Choice Questions 725Answers to Quantitative & Multiple-Choice Questions 775Index 781Tables

Standard Atomic Weights (1995) Inside Front CoverRanges of Normal Values in Human Whole Blood, Plasma, or Serum Inside Back CoverPrefaceThis book is designed to provide a concise summary of mammalian and, particularly, of humanphysiology that medical students and others can use by itself or can supplement with readings in currenttexts, monographs, and reviews. Pertinent aspects of general and comparative physiology are alsoincluded. Summaries of relevant anatomic considerations will be found in each section, but this book iswritten primarily for those who have some knowledge of anatomy, chemistry, and biochemistry. Examplesfrom clinical medicine are given where pertinent to illustrate physiologic points. In many of the chapters,physicians desiring to use this book as a review will find short discussions of important symptomsproduced by disordered function.Review of Medical Physiology also includes a self-study section to help students review for Board andother examinations and an appendix that contains general references, a discussion of statistical methods,a glossary of abbreviations, acronyms, and symbols commonly used in physiology, and several usefultables. The index is comprehensive and specifically designed for ease in locating important terms, topics,and concepts.In writing this book, the author has not been able to be complete and concise without also beingdogmatic. I believe, however, that the conclusions presented without detailed discussion of theexperimental data on which they are based are supported by the bulk of the currently available evidence.Much of this evidence can be found in the papers cited in the credit lines accompanying the illustrations.Further discussions of particular subjects and information on subjects not considered in detail can befound in the references listed at the end of each section. Information about serial review publications thatprovide up-to-date discussions of various physiologic subjects is included in the note on generalreferences in the appendix. In the interest of brevity and clarity, I have in most instances omitted thenames of the many investigators whose work made possible the views of physiology presented here. Thisomission is in no way intended to slight their contributions, but including their names and specificreferences to original papers would greatly increase the length of the book.In this twentieth edition, as in previous editions, the entire book has been thoroughly revised, with a viewto eliminating errors, incorporating suggestions of readers, updating concepts, and discarding materialthat is no longer relevant. In this way, the book has been kept as up-to-date and accurate as possible.Since the last edition, there has continued to be rapid expansion of knowledge about how extracellularsignals initiate changes in gene expression and about the genetic basis of disease. Material on thesetopics has been updated. The section on immunology has been rewritten again for clarity and to expandconsideration of the relation between innate and acquired immunity. The sections on the cerebral cortexin relation to vision, audition, and olfaction have been revised, and the chapter on sleep and wakingstates has been rewritten to emphasize the importance of thalamocortical oscillations. New informationhas been provided on many topics, including molecular motors, hormones of the heart, motilin andgastrointestinal motility, acute phase proteins, sleep apnea, and addiction.The self-study section has been updated, and more emphasis has been placed on physiology in relationto disease, in keeping with the current trend in the United States Medical Licensing Examinations(USMLE).I am greatly indebted to the many individuals who helped with the preparation of this book. Those towhom I express special thanks for their help with the twentieth edition include Dr. Walter Miller, Dr. MelvinGrumbach, Dr. Stephen McPhee, and Dr. Dolores Shoback. Jesse Loesberg provided invaluablesecretarial assistance, and, as always, my wife made numerous contributions. Jim Ransom, who editedthe first edition of this book 40 years ago, came back again and did an excellent job of editing this edition.Many associates and friends provided unpublished illustrative materials, and numerous authors andpublishers generously granted permission to reproduce illustrations from other books and journals. I alsothank all the students and others who took the time to write to me offering helpful criticisms andsuggestions. Such comments are always welcome, and I solicit additional corrections and criticisms,which may be addressed to me at

Department of PhysiologyUniversity of CaliforniaSan Francisco, CA 94143-0444, USASince this book was first published in 1963, the following translations have been published: Bulgarian,Chinese (two independent translations), Czech (two editions), French, German (four editions), Greek (twoeditions), Hungarian, Indonesian (three editions), Italian (seven editions), Japanese (fifteen editions),Korean, Malaysian, Polish (two editions), Portuguese (seven editions), Serbo-Croatian, Spanish (sixteeneditions), and Turkis

Bone Physiology 370 Vitamin D & the Hydroxycholecalciferols 375 The Parathyroid Glands 377 Calcitonin 380 Effects of Other Hormones & Humoral Agents on Calcium Metabolism 382 22. The Pituitary Gland 383 Introduction 383 Morphology 381 Intermediate-Lobe Hormones 384 Growth Hormone 386 Phys

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