Textbook of Medical Physiology.pdf

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1、 TEXTBOOK of Medical Physiology TEXTBOOK of Medical Physiology E L E V E N T H E D I T I O N Arthur C Guyton M D Professor Emeritus Department of Physiology and Biophysics University of Mississippi Medical Center Jackson Mississippi Deceased John E Hall Ph D Professor and Chairman Department of Phys

2、iology and Biophysics University of Mississippi Medical Center Jackson Mississippi Elsevier Inc 1600 John F Kennedy Blvd Suite 1800 Philadelphia Pennsylvania 19103 2899 TEXTBOOK OF MEDICAL PHYSIOLOGYISBN 0 7216 0240 1 International Edition ISBN 0 8089 2317 X Copyright 2006 2000 1996 1991 1986 1981 1

3、976 1971 1966 1961 1956 by Elsevier Inc All rights reserved No part of this publication may be reproduced or transmitted in any form or by any means electronic or mechanical including photocopying recording or any information storage and retrieval system without permission in writing from the publis

4、her Permissions may be sought directly from Elsevier s Health Sciences Rights Department in Philadelphia PA USA phone 1 215 239 3804 fax 1 215 239 3805 e mail healthpermissions You may also complete your request on line via the Elsevier homepage by selecting Customer Support and then Obtaining Permi

5、ssions NOTICE Knowledge and best practice in this fi eld are constantly changing As new research and experience broaden our knowledge changes in practice treatment and drug therapy may become necessary or appropriate Readers are advised to check the most current information provided i on procedures

6、featured or ii by the manufacturer of each product to be administered to verify the recommended dose or formula the method and duration of administration and contraindications It is the responsibility of the practitioner relying on their own experience and knowledge of the patient to make diagnoses

7、to determine dosages and the best treatment for each individual patient and to take all appropriate safety precautions To the fullest extent of the law neither the Publisher nor the Author assumes any liability for any injury and or damage to persons or property arising out or related to any use of

8、the material contained in this book Library of Congress Cataloging in Publication Data Guyton Arthur C Textbook of medical physiology Arthur C Guyton John E Hall 11th ed p cm Includes bibliographical references and index ISBN 0 7216 0240 1 1 Human physiology 2 Physiology Pathological I Title Medical

9、 physiology II Hall John E John Edward III Title DNLM 1 Physiological Processes QT 104 G992t 2006 QP34 5 G92006 612 dc22 2004051421 Publishing Director Linda Belfus Acquisitions Editor William Schmitt Managing Editor Rebecca Gruliow Publishing Services Manager Tina Rebane Project Manager Mary Anne F

10、olcher Design Manager Steven Stave Marketing Manager John Gore Cover illustration is a detail from Opus 1972 by Virgil Cantini Ph D with permission of the artist and Mansfi eld State College Mansfi eld Pennsylvania Chapter opener credits Chapter 43 modifi ed from Getty Images 21000058038 Chapter 44

11、modifi ed from Getty Images 21000044598 Chapter 84 modifi ed from Corbis Printed in China Last digit is the print number 987654321 Working together to grow libraries in developing countries www bookaid org www sabre org To My Family For their abundant support for their patience and understanding and

12、 for their love To Arthur C Guyton For his imaginative and innovative research For his dedication to education For showing us the excitement and joy of physiology And for serving as an inspirational role model Arthur C Guyton M D 1919 2003 I N M E M O R I A M The sudden loss of Dr Arthur C Guyton in

13、 an automobile accident on April 3 2003 stunned and saddened all who were privileged to know him Arthur Guyton was a giant in the fi elds of physiology and medicine a leader among leaders a master teacher and an inspiring role model throughout the world Arthur Clifton Guyton was born in Oxford Missi

14、ssippi to Dr Billy S Guyton a highly respected eye ear nose and throat specialist who later became Dean of the University of Mississippi Medical School and Kate Small wood Guyton a mathematics and physics teacher who had been a missionary in China before marriage During his formative years Arthur en

15、joyed watching his father work at the Guyton Clinic playing chess and swapping stories with William Faulkner and building sailboats one of which he later sold to Faulkner He also built countless mechanical and electrical devices which he continued to do throughout his life His brilliance shone early

16、 as he graduated top in his class at the University of Mississippi He later distinguished himself at Harvard Medical School and began his postgraduate surgical training at Massachusetts General Hospital His medical training was interrupted twice once to serve in the Navy during World War II and agai

17、n in 1946 when he was stricken with poliomyelitis during his fi nal year of residency training Suffering paralysis in his right leg left arm and both shoulders he spent nine months in Warm Springs Georgia recuper ating and applying his inventive mind to building the fi rst motorized wheelchair contr

18、olled by a joy stick a motorized hoist for lifting patients special leg braces and other devices to aid the handicapped For those inventions he received a Presidential Citation He returned to Oxford where he devoted himself to teaching and research at the University of Mississippi School of Medicine

19、 and was named Chair of the Department of Physiology in 1948 In 1951 he was named one of the ten out standing men in the nation When the University of Mississippi moved its Medical School to Jackson in 1955 he rapidly developed one of the world s premier cardiovascular research programs His remarkab

20、le life as a scientist author and devoted father is detailed in a biography published on the occasion of his retirement in 1989 1 A Great Physiologist Arthur Guyton s research contributions which include more than 600 papers and 40 books are legendary and place him among the greatest physiologists i

21、n history His research covered virtually all areas of car diovascular regulation and led to many seminal concepts that are now an inte gral part of our understanding of cardiovascular disorders such as hypertension heart failure and edema It is diffi cult to discuss cardiovascular physiology without

22、 including his concepts of cardiac output and venous return negative interstitial fl uid pressure and regulation of tissue fl uid volume and edema regulation of tissue blood fl ow and whole body blood fl ow autoregulation renal pressure natriuresis and long term blood pressure regulation Indeed his

23、concepts of cardiovascular regulation are found in virtually every major text book of physiology They have become so familiar that their origin is sometimes forgotten One of Dr Guyton s most important scientifi c legacies was his application of principles of engineering and systems analysis to cardi

24、ovascular regulation He used mathematical and graphical methods to quantify various aspects of circu latory function before computers were widely available He built analog com puters and pioneered the application of large scale systems analysis to modeling the cardiovascular system before the advent

25、 of digital computers As digital computers became available his cardiovascular models expanded dramatically to include the kidneys and body fl uids hormones and the autonomic nervous system as well as cardiac and circulatory functions 2 He also provided the fi rst comprehensive systems analysis of b

26、lood pressure regulation This unique approach to physiological research preceded the emergence of biomedical vii viiiIn Memoriam engineering a fi eld that he helped to establish and to promote in physiology leading the discipline into a quantitative rather than a descriptive science It is a tribute

27、to Arthur Guyton s genius that his concepts of cardiovascular regulation often seemed heretical when they were fi rst presented yet stimu lated investigators throughout the world to test them experimentally They are now widely accepted In fact many of his concepts of cardiovascular regulation are in

28、tegral components of what is now taught in most medical physiology courses They continue to be the foundation for generations of cardiovascular physiologists Dr Guyton received more than 80 major honors from diverse scientifi c and civic organizations and uni versities throughout the world A few of

29、these that are especially relevant to cardiovascular research include the Wiggers Award of the American Physiological Society the Ciba Award from the Council for High Blood Pressure Research The William Harvey Award from the American Society of Hypertension the Research Achievement Award of the Amer

30、ican Heart Association and the Merck Sharp so too is feed back important for progressive improvement of a textbook of physiology To the many persons who have already helped I send sincere thanks xi xiiPreface A brief explanation is needed about several features of the 11th edition Although many of t

31、he chapters have been revised to include new principles of physi ology the text length has been closely monitored to limit the book size so that it can be used effec tively in physiology courses for medical students and health care professionals Many of the fi gures have also been redrawn and are no

32、w in full color New references have been chosen primarily for their pres entation of physiologic principles for the quality of their own references and for their easy accessibility Most of the selected references are from recently published scientifi c journals that can be freely accessed from the P

33、ubMed internet site at http www ncbi nlm nih gov entrez query fcgi db PubMed Use of these references as well as cross references from them can give the student almost complete cov erage of the entire fi eld of physiology Another feature is that the print is set in two sizes The material in small pri

34、nt is of several different kinds fi rst anatomical chemical and other information that is needed for immediate discussion but that most stu dents will learn in more detail in other courses second physiologic information of special importance to certain fi elds of clinical medicine and third informa

35、tion that will be of value to those students who may wish to study particular physiologic mechanisms more deeply The material in large print constitutes the funda mental physiologic information that students will require in virtually all their medical activities and studies I wish to express my than

36、ks to many other persons who have helped in preparing this book including my colleagues in the Department of Physiology The Heart as a Pump and Function of the Heart Valves103 Physiology of Cardiac Muscle103 Physiologic Anatomy of Cardiac Muscle103 Action Potentials in Cardiac Muscle104 The Cardiac

37、Cycle106 Diastole and Systole106 Relationship of the Electrocardiogram to the Cardiac Cycle107 Function of the Atria as Primer Pumps107 Function of the Ventricles as Pumps108 Table of Contentsxv Function of the Valves109 Aortic Pressure Curve109 Relationship of the Heart Sounds to Heart Pumping109 W

38、ork Output of the Heart110 Graphical Analysis of Ventricular Pumping110 Chemical Energy Required for Cardiac Contraction Oxygen Utilization by the Heart111 Regulation of Heart Pumping111 Intrinsic Regulation of Heart Pumping The Frank Starling Mechanism111 Effect of Potassium and Calcium Ions on Hea

39、rt Function113 Effect of Temperature on Heart Function114 Increasing the Arterial Pressure Load up to a Limit Does Not Decrease the Cardiac Output114 C H A P T E R 1 0 Rhythmical Excitation of the Heart116 Specialized Excitatory and Conductive System of the Heart116 Sinus Sinoatrial Node116 Internod

40、al Pathways and Transmission of the Cardiac Impulse Through the Atria118 Atrioventricular Node and Delay of Impulse Conduction from the Atria to the Ventricles118 Rapid Transmission in the Ventricular Purkinje System119 Transmission of the Cardiac Impulse in the Ventricular Muscle119 Summary of the

41、Spread of the Cardiac Impulse Through the Heart120 Control of Excitation and Conduction in the Heart120 The Sinus Node as the Pacemaker of the Heart120 Role of the Purkinje System in Causing Synchronous Contraction of the Ventricular Muscle121 Control of Heart Rhythmicity and Impulse Conduction by t

42、he Cardiac Nerves The Sympathetic and Parasympathetic Nerves121 C H A P T E R 1 1 The Normal Electrocardiogram123 Characteristics of the Normal Electrocardiogram123 Depolarization Waves Versus Repolarization Waves123 Relationship of Atrial and Ventricular Contraction to the Waves of the Electrocardi

43、ogram125 Voltage and Time Calibration of the Electrocardiogram125 Methods for Recording Electrocardiograms126 Pen Recorder126 Flow of Current Around the Heart During the Cardiac Cycle126 Recording Electrical Potentials from a Partially Depolarized Mass of Syncytial Cardiac Muscle126 Flow of Electric

44、al Currents in the Chest Around the Heart126 Electrocardiographic Leads127 Three Bipolar Limb Leads127 Chest Leads Precordial Leads 129 Augmented Unipolar Limb Leads129 C H A P T E R 1 2 Electrocardiographic Interpretation of Cardiac Muscle and Coronary Blood Flow Abnormalities Vectorial Analysis131

45、 Principles of Vectorial Analysis of Electrocardiograms131 Use of Vectors to Represent Electrical Potentials131 Direction of a Vector Is Denoted in Terms of Degrees131 Axis for Each Standard Bipolar Lead and Each Unipolar Limb Lead132 Vectorial Analysis of Potentials Recorded in Different Leads133 V

46、ectorial Analysis of the Normal Electrocardiogram134 Vectors That Occur at Successive Intervals During Depolarization of the Ventricles The QRS Complex134 Electrocardiogram During Repolarization The T Wave134 Depolarization of the Atria The P Wave136 Vectorcardiogram136 Mean Electrical Axis of the V

47、entricular QRS And Its Signifi cance137 Determining the Electrical Axis from Standard Lead Electrocardiograms137 Abnormal Ventricular Conditions That Cause Axis Deviation138 Conditions That Cause Abnormal Voltages of the QRS Complex140 Increased Voltage in the Standard Bipolar Limb Leads140 Decrease

48、d Voltage of the Electrocardiogram140 Prolonged and Bizarre Patterns of the QRS Complex141 Prolonged QRS Complex as a Result of Cardiac Hypertrophy or Dilatation141 Prolonged QRS Complex Resulting from Purkinje System Blocks141 Conditions That Cause Bizarre QRS Complexes141 Current of Injury141 Effe

49、ct of Current of Injury on the QRS Complex141 The J Point The Zero Reference Potential for Analyzing Current of Injury142 Coronary Ischemia as a Cause of Injury Potential143 Abnormalities in the T Wave145 Effect of Slow Conduction of the Depolarization Wave on the Characteristics of the T Wave145 Sh

50、ortened Depolarization in Portions of the Ventricular Muscle as a Cause of T Wave Abnormalities145 xviTable of Contents C H A P T E R 1 3 Cardiac Arrhythmias and Their Electrocardiographic Interpretation147 Abnormal Sinus Rhythms147 Tachycardia147 Bradycardia147 Sinus Arrhythmia148 Abnormal Rhythms