T E X T B K 0 F EISEVTER Medical e Physiology E L E V E N TH E I0 N purposes only GUYToN HALL
T E X T 0 K of medical Physiology E LE VENTH EDTION Arthur C. Guyton, M.D.t Professor emeritus Department of physiology and biophysics University of Mississippi Medical Center Jackson, Mississippi Deceased John e hall ph D Professor and Chairman Department of Physiology and Biophysics University of Mississippi Medical Center Jackson, Mississippi ELSEVIER SAUNDERS
TEXTBOOK of Medical Physiology ELEVENTH EDITION Arthur C. Guyton, M.D.† Professor Emeritus Department of Physiology and Biophysics University of Mississippi Medical Center Jackson, Mississippi † Deceased John E. Hall, Ph.D. Professor and Chairman Department of Physiology and Biophysics University of Mississippi Medical Center Jackson, Mississippi
ELSEVIER SAUNDERS Elsevier lnc. 1600 John F Kennedy Blvd. Suite 1800 Philadelphia, Pennsylvania 19103-2899 TEXTBOOK OF MEDICAL PHYSIOLOGY ISBN0-7216-0240-1 International Edition Isbn 0-8089-2317-X 6,1991,1986,1981,1976,1971,196,1961,1956 by elsevier Inc. All rights reserved. No part of this publication duced or transmitted in any form or by any means, electronic or mechanical, including ng, recording, or any information stor publisher. Permissions may be sought Philadelphia, PA, USA: Phone: (+1) 152393804,fax(+1) 9 3805.e-mail: healt @elsevier.com.Youmayalsocomplete yourrequeston-lineviatheElsevierhomepage(http://www.elsevier.com),byselecting"custome Support"and then"Obtaining Permissions NOTICE our knowledge, changes in practice, treatment and drug therapy may become experienc Ige and best practice in this field are constantly changing. As new research an essary or appropriate. Readers are advised to check the most current information provided (i)on ocedures 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. g eIr patient, to make diagnoses, to determine dosages and the best treatment for each individual tient, 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 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-1lth ed. Includes bibliographical references and index. ISBN07216-0240-1 DNLM: 1. Physiological Processes. QT 104 G992t 2006 oP34.5G92006 612dc22 004051421 Publishing Director: Linda belfus Managing Editor: Rebecca Gruliow Publishing Services Mi Tina Rebane Project Manager: Mary Anne Folcher 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 Mansfield State College. Mansfield, Pennsylvania Chapter opener credits: Chapter 43, modified from Getty Images 21000058038: Chapter 44, modified from Getty Images 21000044598: Chapter 84, modified from Corbis ng toget grow libraries in developing countries Printed in China www.elsevicr.comIwww.bookaid.orgIwww.sabre.org Last digit is the umber:98765432 El SEVIER BOOK AID Sabre Foundati
Elsevier Inc. 1600 John F. Kennedy Blvd., Suite 1800 Philadelphia, Pennsylvania 19103-2899 TEXTBOOK OF MEDICAL PHYSIOLOGY ISBN 0-7216-0240-1 International Edition ISBN 0-8089-2317-X Copyright © 2006, 2000, 1996, 1991, 1986, 1981, 1976, 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 publisher. 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@elsevier.com. You may also complete your request on-line via the Elsevier homepage (http://www.elsevier.com), by selecting “Customer Support” and then “Obtaining Permissions”. NOTICE Knowledge and best practice in this field 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 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, 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 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 physiology. II. Hall, John E. (John Edward) III. Title. [DNLM: 1. Physiological Processes. QT 104 G992t 2006] QP34.5.G9 2006 612—dc22 2004051421 Publishing Director: Linda Belfus Acquisitions Editor: William Schmitt Managing Editor: Rebecca Gruliow Publishing Services Manager: Tina Rebane Project Manager: Mary Anne Folcher 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 Mansfield State College, Mansfield, Pennsylvania. Chapter opener credits: Chapter 43, modified from © Getty Images 21000058038; Chapter 44, modified from © Getty Images 21000044598; Chapter 84, modified from © Corbis. Printed in China Last digit is the print number: 987654321 Working together to grow libraries in developing countries www.elsevier.com | www.bookaid.org | www.sabre.org
MY FAMILY For their abundant support, for their patience and understanding, and for their love 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
To My Family For their abundant support, for their patience and understanding, and for their love To Arthur C. Guyton For his imaginative and innovative research For his dedication to education For showing us the excitement and joy of physiology And for serving as an inspirational role model
Arthur C. Guyton, MD 1919-2003
Arthur C. Guyton, M.D. 1919–2003
IN MEMORIAM The sudden loss of Dr Arthur C Guyton in an automobile accident on April 3, 2003, stunned and saddened all who were privileged to know him. Arthur Guyton was a giant in the fields of physiology and medicine, a leader among leaders, a master teacher, and an inspiring role model throughout the world. Arthur Clifton Guyton was born in Oxford, Mississippi, to Dr. Billy S Guyton, a highly respected eye, ear, nose, and throat specialist, who later became Dean of the University of Mississippi Medical School, and Kate Small in China before marriage. During his formative years, Arthur enjoyed watchin his father work at the Guyton Clinic, playing chess and swapping stories with William Faulkner, and building sailboats (one of which he later sold to Faulkner ). He also built countless mechanical and electrical devices, which he continued to do throughout his life. His brilliance shone early as he graduated top in his class at the University of Mississippi. He later distinguished himself at Harvard Medical School and began his postgraduate surgical training at Massachusetts General Hospital His medical training was interrupted twice-once to serve in the Navy during World War II and again in 1946 when he was str ricken with poliomyelitis during his final 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 first motorized wheelchair controlled by a"joy stick, "a motorized hoist for lifting patients, special leg braces, and other devices to aid the handicapped. For those inventions he received a Presidential Citation He returned to Oxford where he devoted himself to teaching and research at the University of Mississippi School of medicine and was named Chair of the Department of Physiology in 1948. In 1951 he was named one of the ten out standing men in the nation. When the University of Mississippi moved its Medical School to Jackson in 1955, he rapidly developed one of the world's premier cardiovascular research programs. His remarkable life as a scientist author, and devoted father is detailed in a biography published on the occasion of his“ retirement”in1989 A Great Physiologist. Arthur Guyton's research contributions, which include more than 600 papers and 40 books, are legendary and place him among the eatest physiologists in history. His research covered virtually all areas of car diovascular regulation and led to many seminal concepts that are now an inte gral part of our understanding of cardiovascular disorders, such as hypertension heart failure, and edema. It is difficult to discuss cardiovascular physiology without including his concepts of cardiac output and venous return, negative interstitial fluid pressure and regulation of tissue fiuid volume and edema regulation of tissue blood flow and whole body blood flow autoregulation renal-pressure natriuresis, and long-term blood pressure regulation. Indeed, his concepts of cardiovascular regulation are found in virtually every major text book of physiology. They have become so familiar that their origin is sometimes forgotten One of Dr Guyton,s most important scientific legacies was his application of principles of engineering and systems analysis to cardiovascular regulation. He used mathematical and graphical methods to quantify various aspects of circu puters axa Pioneered the application of large-scale systems analysis to modeling latory function before computers were widely available. He built analog cor the cardiovascular system before the advent of digital computers. As digital computers became available, his cardiovascular models expanded dramatically to include the kidneys and body fluids, hormones, and the autonomic nervous system, as well as cardiac and circulatory functions. He also provided the first comprehensive systems analysis of blood pressure regulation. This unique approach to physiological research preceded the emergence of biomedical
IN MEMORIAM The sudden loss of Dr. Arthur C. Guyton in an automobile accident on April 3, 2003, stunned and saddened all who were privileged to know him. Arthur Guyton was a giant in the fields of physiology and medicine, a leader among leaders, a master teacher, and an inspiring role model throughout the world. Arthur Clifton Guyton was born in Oxford, Mississippi, to Dr. Billy S. Guyton, a highly respected eye, ear, nose, and throat specialist, who later became Dean of the University of Mississippi Medical School, and Kate Smallwood Guyton, a mathematics and physics teacher who had been a missionary in China before marriage. During his formative years, Arthur enjoyed watching his father work at the Guyton Clinic, playing chess and swapping stories with William Faulkner, and building sailboats (one of which he later sold to Faulkner). He also built countless mechanical and electrical devices, which he continued to do throughout his life. His brilliance shone early as he graduated top in his class at the University of Mississippi. He later distinguished himself at Harvard Medical School and began his postgraduate surgical training at Massachusetts General Hospital. His medical training was interrupted twice—once to serve in the Navy during World War II and again in 1946 when he was stricken with poliomyelitis during his final year of residency training. Suffering paralysis in his right leg, left arm, and both shoulders, he spent nine months in Warm Springs, Georgia, recuperating and applying his inventive mind to building the first motorized wheelchair controlled by a “joy stick,” a motorized hoist for lifting patients, special leg braces, and other devices to aid the handicapped. For those inventions he received a Presidential Citation. He returned to Oxford where he devoted himself to teaching and research at the University of Mississippi School of Medicine and was named Chair of the Department of Physiology in 1948. In 1951 he was named one of the ten outstanding 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 remarkable life as a scientist, author, and devoted father is detailed in a biography published on the occasion of his “retirement” in 1989.1 A Great Physiologist. Arthur Guyton’s research contributions, which include more than 600 papers and 40 books, are legendary and place him among the greatest physiologists in history. His research covered virtually all areas of cardiovascular regulation and led to many seminal concepts that are now an integral part of our understanding of cardiovascular disorders, such as hypertension, heart failure, and edema. It is difficult to discuss cardiovascular physiology without including his concepts of cardiac output and venous return, negative interstitial fluid pressure and regulation of tissue fluid volume and edema, regulation of tissue blood flow and whole body blood flow autoregulation, renal-pressure natriuresis, and long-term blood pressure regulation. Indeed, his concepts of cardiovascular regulation are found in virtually every major textbook of physiology.They have become so familiar that their origin is sometimes forgotten. One of Dr. Guyton’s most important scientific legacies was his application of principles of engineering and systems analysis to cardiovascular regulation. He used mathematical and graphical methods to quantify various aspects of circulatory function before computers were widely available. He built analog computers and pioneered the application of large-scale systems analysis to modeling the cardiovascular system before the advent of digital computers. As digital computers became available, his cardiovascular models expanded dramatically to include the kidneys and body fluids, hormones, and the autonomic nervous system, as well as cardiac and circulatory functions.2 He also provided the first comprehensive systems analysis of blood pressure regulation. This unique approach to physiological research preceded the emergence of biomedical vii
In memoriam engineering-a field that he helped to and to and 20/20 described the remarkable home environ promote in physiology, leading the ito a ment that Arthur and Ruth Guyton created to raise quantitative rather than a descriptive their family. His devotion to family is beautifully It is a tribute to Arthur Guyton's genius that his expressed in the dedication of his Textbook of medical concepts of cardiovascular regulation often seemed Physiology heretical when they were first presented, yet stimu- lated investigators throughout the world to test them experimentally. They are now widely accepted. In fact, My father for his uncompromising principles that many of his concepts of cardiovascular regulation uided my life are integral components of what is now taught in My mother for leading her children into intellectual most medical physiology courses. They continue to pursuits be the foundation for generations of cardiovascular My wife for her magnificent devotion to her family My children for making everything worthwhile Dr. Guyton received more than 80 major honors Dr Guyton was a master teacher at the Universit from diverse scientific and civic organizations and uni- of Mississippi for over 50 years. Even though he was versities throughout the world. A few of these that are always busy with service responsibilities, research especially relevant to cardiovascular research include writing, and teaching, he was never too busy to talk the Wiggers Award of the American Physiological with a student who was having difficulty. He would Blood Pressure Research, The William Harvey Award if it conflicted with his teaching schedule gious lecture Society, the Ciba Award from the Council for High never accept an invitation to give a prestig from the American Society of Hypertension, the His contributions to education are also far reach Research Achievement Award of the American Heart ing through generations of physiology graduate Association, and the Merck Sharp Dohme Award students and postdoctoral fellows. He trained over of the International Society of Hypertension. It was 150 scientists, at least 29 of whom became chairs of appropriate that in 1978 he was invited by the Royal their own departments and six of whom became pres- College of Physicians in London to deliver a special idents of the American Physiological Society. He gave lecture honoring the 400th anniversary of the birth of students confidence in their abilities and emphasized william Harvey, who discovered the circulation of the his belief that "People who are really successful in the blood research world are self-taught. He insisted that his Dr. Guyton's love of physiology was beautifully trainees integrate their experimental findings into a articulated in his presidents address to the American broad conceptual framework that included other Physiological Society in 1975, appropriately entitled interacting systems. This approach usually led them Physiology, a Beauty and a Philosophy. Let me quote to develop a quantitative analysis and a better just one sentence from his address: What other person, understanding of the particular physiological systems whether he be a theologian, a jurist, a doctor of medi that they were studying. No one has been more pro- ine, a physicist, or whatever, knows more than you, a lific in training leaders of physiology than Arthur physiologist, about life? For physiology is indeed an Guyton explanation of life. What other subject matter is more Dr Guyton,s Textbook of Medical Physiology, first fascinating, more exciting, more beautiful than the published in 1956, quickly became the best-selling subject of life? medical physiology textbook in the world. He had a ft for communicating complex ideas in a clear and A Master Teacher. Although Dr. Guyton's accomplishments are legendary, his contribution ducator have probably had an even greater He and his wonderful wife ruth raised ten m interesting manner that made studying physiology fun He wrote the book to teach his students, not to impress his professional colleagues. Its popularity with stu- dents has made it the most widely used physiology all of whom became outstanding physicians-a textbook in history. This accomplishment alone was remarkable educational achievement. Eight of the enough to ensure his legacy Guyton children graduated from Harvard Medical The Textbook of Medical Physiology began as School, one from duke medical school and one from lecture notes in the early 1950s when Dr Guyton was The University of Miami Medical School after receiv- teaching the entire physiology course for medical stu- ng a Ph.D. from Harvard. An article published in dents at the University of Mississippi. He discovered Reader's Digest in 1982 highlighted their extraordinary that the students were having difficulty with the text family life. books that were available and beg distributing The success of the Guyton children did not occur by copies of his lecture notes. In describing his experi chance. Dr Guyton,s philosophy of education was to ence, Dr. Guyton stated that "Many textbooks of learn by doing. "The children participated in count- medical physiology had become discursive, written pri less family projects that included the design and marily by teachers of physiology for other teachers of construction of their home and its heating system, physiology, and written in language understood by the swimming pool, tennis court, sailboats, go-carts other teachers but not easily understood by the basic and electrical cars, household gadgets, and electronic student of medical physiology instruments for their Oxford Instruments Company. Through his Textbook of Medical Physiology, which Television programs such as Good Morning America is translated into 13 languages, he has probably done
viii In Memoriam engineering—a field that he helped to establish and to promote in physiology, leading the discipline into a quantitative rather than a descriptive science. It is a tribute to Arthur Guyton’s genius that his concepts of cardiovascular regulation often seemed heretical when they were first presented, yet stimulated investigators throughout the world to test them experimentally. They are now widely accepted. In fact, many of his concepts of cardiovascular regulation are integral components of what is now taught in most medical physiology courses. They continue to be the foundation for generations of cardiovascular physiologists. Dr. Guyton received more than 80 major honors from diverse scientific and civic organizations and universities throughout the world. A few of these that are especially relevant to cardiovascular research include the Wiggers Award of the American Physiological Society, the Ciba Award from the Council for High Blood Pressure Research, The William Harvey Award from the American Society of Hypertension, the Research Achievement Award of the American Heart Association, and the Merck Sharp & Dohme Award of the International Society of Hypertension. It was appropriate that in 1978 he was invited by the Royal College of Physicians in London to deliver a special lecture honoring the 400th anniversary of the birth of William Harvey, who discovered the circulation of the blood. Dr. Guyton’s love of physiology was beautifully articulated in his president’s address to the American Physiological Society in 1975,3 appropriately entitled Physiology, a Beauty and a Philosophy. Let me quote just one sentence from his address: What other person, whether he be a theologian, a jurist, a doctor of medicine, a physicist, or whatever, knows more than you, a physiologist, about life? For physiology is indeed an explanation of life. What other subject matter is more fascinating, more exciting, more beautiful than the subject of life? A Master Teacher. Although Dr. Guyton’s research accomplishments are legendary, his contributions as an educator have probably had an even greater impact. He and his wonderful wife Ruth raised ten children, all of whom became outstanding physicians—a remarkable educational achievement. Eight of the Guyton children graduated from Harvard Medical School, one from Duke Medical School, and one from The University of Miami Medical School after receiving a Ph.D. from Harvard. An article published in Reader’s Digest in 1982 highlighted their extraordinary family life.4 The success of the Guyton children did not occur by chance. Dr. Guyton’s philosophy of education was to “learn by doing.” The children participated in countless family projects that included the design and construction of their home and its heating system, the swimming pool, tennis court, sailboats, go-carts and electrical cars, household gadgets, and electronic instruments for their Oxford Instruments Company. Television programs such as Good Morning America and 20/20 described the remarkable home environment that Arthur and Ruth Guyton created to raise their family. His devotion to family is beautifully expressed in the dedication of his Textbook of Medical Physiology5 : To My father for his uncompromising principles that guided my life My mother for leading her children into intellectual pursuits My wife for her magnificent devotion to her family My children for making everything worthwhile Dr. Guyton was a master teacher at the University of Mississippi for over 50 years. Even though he was always busy with service responsibilities, research, writing, and teaching, he was never too busy to talk with a student who was having difficulty. He would never accept an invitation to give a prestigious lecture if it conflicted with his teaching schedule. His contributions to education are also far reaching through generations of physiology graduate students and postdoctoral fellows. He trained over 150 scientists, at least 29 of whom became chairs of their own departments and six of whom became presidents of the American Physiological Society. He gave students confidence in their abilities and emphasized his belief that “People who are really successful in the research world are self-taught.” He insisted that his trainees integrate their experimental findings into a broad conceptual framework that included other interacting systems. This approach usually led them to develop a quantitative analysis and a better understanding of the particular physiological systems that they were studying. No one has been more prolific in training leaders of physiology than Arthur Guyton. Dr. Guyton’s Textbook of Medical Physiology, first published in 1956, quickly became the best-selling medical physiology textbook in the world. He had a gift for communicating complex ideas in a clear and interesting manner that made studying physiology fun. He wrote the book to teach his students, not to impress his professional colleagues. Its popularity with students has made it the most widely used physiology textbook in history. This accomplishment alone was enough to ensure his legacy. The Textbook of Medical Physiology began as lecture notes in the early 1950s when Dr. Guyton was teaching the entire physiology course for medical students at the University of Mississippi. He discovered that the students were having difficulty with the textbooks that were available and began distributing copies of his lecture notes. In describing his experience, Dr. Guyton stated that “Many textbooks of medical physiology had become discursive, written primarily by teachers of physiology for other teachers of physiology, and written in language understood by other teachers but not easily understood by the basic student of medical physiology.”6 Through his Textbook of Medical Physiology, which is translated into 13 languages, he has probably done
In memoriam more to teach physiology to the world than any other We celebrate the magnificent life of Arthur Guyton, individual in history. Unlike most major textbook recognizing that we owe him an enormous debt. which often have 20 or more authors, the first eight gave us an imaginative and innovative approach to editions were written entirely by Dr. Guyton-a feat research and many new scientific concepts. He gave hat is unprecedented for any major medical textbook countless students throughout the world a means of For his many contributions to medical education, Dr. understanding physiology and he gave many of us Guyton received the 1996 Abraham Flexner Award exciting research careers. Most of all, he inspired us- from the Association of American Medical Colleges with his devotion to education, his unique ability to AAMC). According to the AAMC, Arthur Guyton bring out the best in those around him, his warm and for the past 50 years has made an unparalleled generous spirit, and his courage. We will miss him impact on medical education. "He is also honored each tremendously, but he will remain in our memories as year by The American Physiological Society through a shining example of the very best in humanity. Arthur he Arthur C Guyton Teaching Award Guyton was a real hero to the world and his legacy is everlasting. An Inspiring Role Model. Dr. Guyton,s accomplish ments extended far beyond science, medicine, and edu References cation. He was an inspiring role model for life as well as for science. No one was more inspirational or influ- 1. Brinson C, Quinn J: Arthur C. Guyton-His Life. His ential on my scientific career than Dr Guyton. He Family, His Achievements. Jackson, MS, Hederman taught his students much more than physiology Brothers Press. 1989 he taught us life, not so much by what he said but by 2. Guyton AC, Coleman TG, Granger HJ: Circulation his unspoken courage and dedication to the highest overall regulation. Ann Rev Physiol 34: 13-46, 1972 standard Guyton AC: Past-President's Address. Physiology, a He had a special ability to motivate people through Beauty and a Philosophy. The Physiologist 8: 495-501 his indomitable spirit. Although he 1975 lenged by polio, those of us who worked with him 4. Bode r:a doctor who's dad to seven doctors -So far! never thought of him as being handicapped. We were Readers'Digest, December, 1982, pp 141-145 5. Guyton AC: Textbook of Medical Physiology. Philadel too busy trying to keep up with him! His brilliant phia, Saunders, 1956. mind. his indefatigable devotion to science. education 6. Guyton AC: An author's philosophy of ph and family, and his spirit captivated students and book writing. Adv Physiol Ed 19: s1-55, 199 trainees, professional colleagues, politicians, business leaders, and virtually everyone who knew him. He JOHN E. HALL would not succumb to the effects of polio. His courage Jackson, MississippI challenged and inspired us. He expected the best and somehow brought out the very best in people
In Memoriam ix more to teach physiology to the world than any other individual in history. Unlike most major textbooks, which often have 20 or more authors, the first eight editions were written entirely by Dr. Guyton—a feat that is unprecedented for any major medical textbook. For his many contributions to medical education, Dr. Guyton received the 1996 Abraham Flexner Award from the Association of American Medical Colleges (AAMC). According to the AAMC, Arthur Guyton “. . . for the past 50 years has made an unparalleled impact on medical education.” He is also honored each year by The American Physiological Society through the Arthur C. Guyton Teaching Award. An Inspiring Role Model. Dr. Guyton’s accomplishments extended far beyond science, medicine, and education. He was an inspiring role model for life as well as for science. No one was more inspirational or influential on my scientific career than Dr. Guyton. He taught his students much more than physiology— he taught us life, not so much by what he said but by his unspoken courage and dedication to the highest standards. He had a special ability to motivate people through his indomitable spirit. Although he was severely challenged by polio, those of us who worked with him never thought of him as being handicapped. We were too busy trying to keep up with him! His brilliant mind, his indefatigable devotion to science, education, and family, and his spirit captivated students and trainees, professional colleagues, politicians, business leaders, and virtually everyone who knew him. He would not succumb to the effects of polio. His courage challenged and inspired us. He expected the best and somehow brought out the very best in people. We celebrate the magnificent life of Arthur Guyton, recognizing that we owe him an enormous debt. He gave us an imaginative and innovative approach to research and many new scientific concepts. He gave countless students throughout the world a means of understanding physiology and he gave many of us exciting research careers. Most of all, he inspired us— with his devotion to education, his unique ability to bring out the best in those around him, his warm and generous spirit, and his courage. We will miss him tremendously, but he will remain in our memories as a shining example of the very best in humanity. Arthur Guyton was a real hero to the world, and his legacy is everlasting. References 1. Brinson C, Quinn J: Arthur C. Guyton—His Life, His Family, His Achievements. Jackson, MS, Hederman Brothers Press, 1989. 2. Guyton AC, Coleman TG, Granger HJ: Circulation: overall regulation. Ann Rev Physiol 34:13–46, 1972. 3. Guyton AC: Past-President’s Address. Physiology, a Beauty and a Philosophy. The Physiologist 8:495–501, 1975. 4. Bode R:A Doctor Who’s Dad to Seven Doctors—So Far! Readers’ Digest, December, 1982, pp. 141–145. 5. Guyton AC: Textbook of Medical Physiology. Philadelphia, Saunders, 1956. 6. Guyton AC: An author’s philosophy of physiology textbook writing. Adv Physiol Ed 19: s1–s5, 1998. John E. Hall Jackson, Mississippi
PR E FA C E The first edition of the Textbook of Medical Phys iology was written by Arthur C Guyton almost 50 years ago. Unlike many major medical textbooks, which often have 20 or more authors. the first eight editions of the Textbook of Medical Physi ology were written entirely by Dr. Guyton with each new edition arriving on schedule for nearly 40 years. Over the years, Dr. Guyton's textbook became widely used throughout the world and was translated into 13 languages A major reason for the book's unprecedented success was his uncanny ability to explain complex physiologic principles in language easily understood by stu- dents. His main goal with each edition was to instruct students in physiology, not to impress his professional colleagues. His writing style always maintained the tone of a teacher talking to his students I had the privilege of working closely with Dr Guyton for almost 30 years nd the honor of helping him with the 9th and 10th editions. For the 1lth edition, I have the same goal as in previous editions--to explain, in language easily understood by students, how the different cells, tissues, and organs of the human body work together to maintain life. This task has been challenging and exciting because our rapidly increasing knowledge of physiology continues to unravel new mysteries of body functions. Many new techniques for learning about molecular and cellular physiology have been developed. We can present more and more the physiology principles in the terminology of molecular and physical sciences rather than in merely a series of separate and unexplained bio- logical phenomena. This change is welcomed, but it also makes revision of each chapter a necessit In this edition, I have attempted to maintain the same unified organization of the text that has been useful to students in the past and to ensure that he book is comprehensive enough that students will wish to use it in later life as a basis for their professional careers. I hope that this textbook conveys the majesty of the human body and its many functions and that it stimulates students to study physiology throughout their careers. Physiology is the link between the basic sciences and medicine. The great beauty of physiology is that it integrates the individual functions of all the body's different cells, tissues, and organs into a functional whole, the human body. Indeed, the human body is much more than the sum of its parts, and life relies upon this total func tion, not just on the function of individual body parts in isolation from the others This brings us to an important question: How are the separate organs and systems coordinated to maintain proper function of the entire body? Fortu nately, our bodies are endowed with a vast network of feedback controls that achieve the necessary balances without which we would not be able to live Physiologists call this high level of internal bodily control homeostasis. I disease states, functional balances are often seriously disturbed and homeosta sis is impaired. And, when even a single disturbance reaches a limit, the whole body can no longer live. One of the goals of this text, therefore, is to emphasize the effectiveness and beauty of the bodys homeostasis mechanisms as well as to present their abnormal function in disease Another objective is to be as accurate as possible. Suggestions and critiques from many physiologists, students, and clinicians throughout the world have been sought and then used to check factual accuracy as well as balance in the text. Even so, because of the likelihood of error in sorting through many thou sands of bits of information, I wish to issue still a further request to all readers to send along notations of error or inaccuracy. Physiologists understand the importance of feedback for proper function of the human body; so, too, is feed the many persons who have already helped, I send sincere thanks siology. To back important for progressive improvement of a textbook of ph
PREFACE The first edition of the Textbook of Medical Physiology was written by Arthur C. Guyton almost 50 years ago. Unlike many major medical textbooks, which often have 20 or more authors, the first eight editions of the Textbook of Medical Physiology were written entirely by Dr. Guyton with each new edition arriving on schedule for nearly 40 years. Over the years, Dr. Guyton’s textbook became widely used throughout the world and was translated into 13 languages. A major reason for the book’s unprecedented success was his uncanny ability to explain complex physiologic principles in language easily understood by students. His main goal with each edition was to instruct students in physiology, not to impress his professional colleagues. His writing style always maintained the tone of a teacher talking to his students. I had the privilege of working closely with Dr. Guyton for almost 30 years and the honor of helping him with the 9th and 10th editions. For the 11th edition, I have the same goal as in previous editions—to explain, in language easily understood by students, how the different cells, tissues, and organs of the human body work together to maintain life. This task has been challenging and exciting because our rapidly increasing knowledge of physiology continues to unravel new mysteries of body functions. Many new techniques for learning about molecular and cellular physiology have been developed. We can present more and more the physiology principles in the terminology of molecular and physical sciences rather than in merely a series of separate and unexplained biological phenomena. This change is welcomed, but it also makes revision of each chapter a necessity. In this edition, I have attempted to maintain the same unified organization of the text that has been useful to students in the past and to ensure that the book is comprehensive enough that students will wish to use it in later life as a basis for their professional careers. I hope that this textbook conveys the majesty of the human body and its many functions and that it stimulates students to study physiology throughout their careers. Physiology is the link between the basic sciences and medicine. The great beauty of physiology is that it integrates the individual functions of all the body’s different cells, tissues, and organs into a functional whole, the human body. Indeed, the human body is much more than the sum of its parts, and life relies upon this total function, not just on the function of individual body parts in isolation from the others. This brings us to an important question: How are the separate organs and systems coordinated to maintain proper function of the entire body? Fortunately, our bodies are endowed with a vast network of feedback controls that achieve the necessary balances without which we would not be able to live. Physiologists call this high level of internal bodily control homeostasis. In disease states, functional balances are often seriously disturbed and homeostasis is impaired. And, when even a single disturbance reaches a limit, the whole body can no longer live. One of the goals of this text, therefore, is to emphasize the effectiveness and beauty of the body’s homeostasis mechanisms as well as to present their abnormal function in disease. Another objective is to be as accurate as possible. Suggestions and critiques from many physiologists, students, and clinicians throughout the world have been sought and then used to check factual accuracy as well as balance in the text. Even so, because of the likelihood of error in sorting through many thousands of bits of information, I wish to issue still a further request to all readers to send along notations of error or inaccuracy. Physiologists understand the importance of feedback for proper function of the human body; so, too, is feedback important for progressive improvement of a textbook of physiology. To the many persons who have already helped, I send sincere thanks. xi
react a brief explanation is needed about several features wish to study particular physiologic mechanisms more of the 11th edition. Although many of the chapters deeply have been revised to include new principles of physi- The material in large print constitutes the funda ology, the text length has been closely monitored mental physiologic information that students will to limit the book size so that it can be used effec- require in virtually all their medical activities and tively in physiology courses for medical students and studies. health care professionals. Many of the figures have I wish to express my thanks to many other persons also been redrawn and are now in full color. New who have helped in preparing this book, includi references have been chosen primarily for their pres- my colleagues in the Department of Physiology entation of physiologic principles, for the quality of Biophysics at the University of Mississippi Medical their own references, and for their easy accessibility. Center who provided valuable suggestions. I am also Most of the selected references are from recently grateful to Ivadelle Osberg Heidke, Gerry McAlpin, published scientific journals that can be freely and Stephanie Lucas for their excellent secretarial accessedfromthePubmedinternetsiteathttp://servicesandtowilliamSchmittRebeccaGruliow www.ncbi.nlm.nihgov/entrezlquery.fcgi?db=pubmed.MaryAnneFolcher,andtherestofthestaffof from them, can give the student almost complete cov- tion excellence s for continued editorial and produc- Use of these references as well as cross-references Elsevier saunder erage of the entire field of physiology Finally, I owe an enormous debt to Arthur Guyton Another feature is that the print is set in two sizes. for an exciting career in physiology, for his friendshi The material in small print is of several different kinds: for the great privilege of contributing to the Textbook is needed for immediate discussion but that most stu- provided to all who knew ha t the inspiration that he first, anatomical, chemical, and other information that of Medical Physiology, and fo dents will learn in more detail in other courses second certain fields of clinical medicine: and. third. informa- JOHN E HALL tion that will be of value to those students who Jackson, Mississippi
xii Preface A brief explanation is needed about several features of the 11th edition. Although many of the chapters have been revised to include new principles of physiology, the text length has been closely monitored to limit the book size so that it can be used effectively in physiology courses for medical students and health care professionals. Many of the figures have also been redrawn and are now in full color. New references have been chosen primarily for their presentation of physiologic principles, for the quality of their own references, and for their easy accessibility. Most of the selected references are from recently published scientific journals that can be freely accessed from the PubMed internet site at http:// www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed. Use of these references, as well as cross-references from them, can give the student almost complete coverage of the entire field of physiology. Another feature is that the print is set in two sizes. The material in small print is of several different kinds: first, anatomical, chemical, and other information that is needed for immediate discussion but that most students will learn in more detail in other courses; second, physiologic information of special importance to certain fields of clinical medicine; and, third, information 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 fundamental physiologic information that students will require in virtually all their medical activities and studies. I wish to express my thanks to many other persons who have helped in preparing this book, including my colleagues in the Department of Physiology & Biophysics at the University of Mississippi Medical Center who provided valuable suggestions. I am also grateful to Ivadelle Osberg Heidke, Gerry McAlpin, and Stephanie Lucas for their excellent secretarial services, and to William Schmitt, Rebecca Gruliow, Mary Anne Folcher, and the rest of the staff of Elsevier Saunders for continued editorial and production excellence. Finally, I owe an enormous debt to Arthur Guyton for an exciting career in physiology, for his friendship, for the great privilege of contributing to the Textbook of Medical Physiology, and for the inspiration that he provided to all who knew him. John E. Hall Jackson, Mississippi
