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GUYTON HALL and and JVlechanisms » lsease ixth edition ^Wk lA^nM^ Human I hysiology and JVlechanisms Disease of Digitized by the Internet Archive 2011 in http://www.archive.org/details/humanphysiologymOOguyt Human Physiology and JVlechanisms ofDi lsease sixth edition MD Arthur C. Guyton, Professor of Physiology and Biophysics Department of Physiology and Biophysics University of Mississippi Medical Center Jackson, Mississippi and John E. Hall, PhD Professor and Chairman Department of Physiology and Biophysics University of Mississippi Medical Center Jackson, Mississippi SAUNDERS COMPANY W.B. A Division ofHarcourt Brace & Company Philadelphia London Toronto Montreal Sydney Tokyo W.B. SAUNDERS COMPANY A Division ofHarcourt Brace &Company TheCurtis Center Independence Square West Philadelphia, Pennsylvania 19106 Library of Congress Cataloging-in-Publication Data Guyton, ArthurC. Human physiology and mechanisms of disease/Arthur C Guyton andJohn E. Hall.—6th ed. p. cm. Includesbibliographical references and index. ISBN 0-7216-3299-8 1. Humanphysiology. 2. Physiology, Pathological. I. Hall,John E. (John Edward). II.Title. [DNLM: 1. Physiology. 2. Disease. QT 104G992b 1997] QP34.5.G87 1997 612—dc20 DNLM/DLC J5-47117 < Portions of this book, including both text and illustrations, have appeared previously in the Text- bookofMedical PhysiologybyArthurC. Guyton andJohn E. Hall, published by W.B. Saunders, 1996. Cover illustration is THE EXPLORER 1970, enamel on steel, by Virgil Cantini, PhD, with permission from the artist, owned by Mr. and Mrs. Richard Natheson, Pittsburgh, PA. HUMAN PHYSIOLOGYAND MECHANISMS OFDISEASE: Sixth Edition 0-7216-3299-8 Copyright © 1997, 1992, 1987, 1982, 1977, 1971 byW.B. Saunders Company. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Printed in the United States ofAmerica. 98765432 Last digit is the print number: Prehace This textbook, entitled Human Physiology and Mechanisms ofDisease, is written for those students who do not have the time to study one of the more formida- ble books and yet require more than the usual college introduction to the physiology of the human body These include medical students, dental students, pharmacy students, other paramedical students in all fields, and many biology students. In general, the types of material presented are threefold: first, the basic fundamental principles of life itself, beginning with the physical, chemical, and molecular laws of cellular function; second, the special concepts and principles necessary for understanding the overall integrated function of the human body; and, third, multiple illustrative examples of how physiological function can be- come abnormal in human disease conditions. This is the sixth edition of this text. The changes from the fifth edition have been many, including update of the physiological understanding of virtually every functional area of the body However, the most extensive changes have come in two special areas: First is the fundamental molecular mechanisms of physiology, a field that is advancing exceedingly rapidly. Fortunately, newer knowledge in this area simplifies many of the explanations for which only speculation could be given previously. The second major area of change has been in the higher functions of the nervous system. Research on nervous mechanisms has always been difficult, so that new fundamental principles of nervous function have been very slow to develop. Yet, within the past decade, especially with the advancing knowledge of neuronal molecular physiology, many of the deeper levels of nervous function are at last beginning to fall in place. We hope especially that this text can convey to the student that our bodies are among the most complex and yet most logical and beautiful of all functional mechanisms. We hope, too, that he or she will understand that each indi- vidual living cell carries within its nucleus all the genetic information required to create an entirely new human being; yet, this same genetic pool serves as al- most 100,000 separate intracellular control systems that regulate the chemical reactions within the cells. And, finally, we hope that all students will recognize, as a special example, the majesty of the human brain as a powerful computer having capabilities that all the electronic computers in the world cannot at present achieve. We could go on detailing the miracles of the human body. That, indeed, is the purpose of this entire text. The success of the book will be measured by the degree of excitement that it leaves with the student for further study in the field of physiology or for a lifetime of physiological thinking. A small but important portion of this text presents not only knowledge that has come from basic experiments in animals but also knowledge that has come from human experiments, especially unplanned experiments caused by disease. For instance, a major share of our knowledge of the regulation of blood glucose and of the mechanisms of carbohydrate metabolism has come from study of di- VI Preface abetes mellitus, a disease that alters these physiological functions profoundly and that is widespread among the human population. Likewise, literally thou- sands of human disease "experiments" proceed each day in the fields of high blood pressure, congestive heart failure, gastrointestinal disturbances, respira- tory diseases, and so forth. The physiology of each of these abnormalities is discussed briefly, partly because study of the diseases themselves can be en- lightening but even more, because they give important insights into basic physiological concepts. Finally, we would like to thank many others who have contributed to the development of this text. For the illustrations we are especially indebted to Tomiko Mita, Michael Schenk, Cathy Garrity, Angela Gardner, and Myriam Kirkman, and for excellent secretarial services we owe our gratitude to Ivadelle Osberg Heidke, Gwendolyn Robbins, and Susie Zuller. We also extend our appreciation to the staff of the W.B. Saunders Company for its continued excel- lence in all publication matters, with appreciation especially to William Schmitt, Editorial Manager; Nicholas Rook, Designer; Linda R. Garber, Senior Produc- tion Manager; Wynette Kommer, Manuscript Editor; and Walter Verbitski, Illus- tration Specialist, whose editorial and technical help have been invaluable. MD Arthur C. Guyton, John E. Hall, PhD