Guyton And Hall Textbook Of Medical Physiology

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Guyton and HallTextbook of Medical .jsp?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatter

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Twelfth EditionGuyton and HallTextbook of Medical PhysiologyJohn E. Hall, Ph.D.Arthur C. Guyton Professor and ChairDepartment of Physiology and BiophysicsAssociate Vice Chancellor for ResearchUniversity of Mississippi Medical CenterJackson, t.jsp?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatter

1600 John F. Kennedy Blvd.Ste 1800Philadelphia, PA 19103-2899TEXTBOOK OF MEDICAL PHYSIOLOGY ISBN: 978-1-4160-4574-8International Edition: 978-0-8089-2400-5Copyright 2011, 2006, 2000, 1996, 1991, 1986, 1981, 1976, 1966,1961, 1956 by Saunders, an imprint of Elsevier Inc.All rights reserved. No part of this publication may be reproduced or transmitted in any formor by any means, electronic or mechanical, including photocopying, recording, or any informationstorage and retrieval system, without permission in writing from the publisher. Permissions may besought directly from Elsevier’s Rights Department: phone: ( 1) 215 239 3804 (US) or ( 44) 1865843830 (UK); fax: ( 44) 1865 853333; e-mail: healthpermissions@elsevier.com. You may alsocomplete your request on-line via the Elsevier website at e and best practice in this field are constantly changing. As new research and experiencebroaden 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 recommendeddose or formula, the method and duration of administration, and contraindications. It is theresponsibility of the practitioner, relying on his or her experience and knowledge of the patient, tomake diagnoses, to determine dosages and the best treatment for each individual patient, and totake all appropriate safety precautions. To the fullest extent of the law, neither the Publisher nor theAuthor assume any liability for any injury and/or damage to persons or property arising out of orrelated to any use of the material contained in this book.The PublisherLibrary of Congress Cataloging-in-Publication DataHall, John E. (John Edward), 1946Guyton and Hall textbook of medical physiology / John Hall. – 12th ed.p. ; cm.Rev. ed. of: Textbook of medical physiology. 11th ed. c2006.Includes bibliographical references and index.ISBN 978-1-4160-4574-8 (alk. paper)1. Human physiology. 2. Physiology, Pathological. I. Guyton, Arthur C. II.Textbook of medical physiology. III. Title. IV. Title: Textbook of medical physiology.[DNLM: 1. Physiological Phenomena. QT 104 H1767g 2011]QP34.5.G9 2011612–dc222009035327Publishing Director: William SchmittDevelopmental Editor: Rebecca GruliowEditorial Assistant: Laura StingelinPublishing Services Manager: Linda Van PeltProject Manager: Frank MoralesDesign Manager: Steve StaveIllustrator: Michael SchenkMarketing Manager: Marla LiebermanPrinted in the United States of AmericaLast digit is the print roduct.jsp?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatter

ToMy FamilyFor their abundant support, for their patience andunderstanding, and for their loveToArthur C. GuytonFor his imaginative and innovative researchFor his dedication to educationFor showing us the excitement and joy of physiologyAnd for serving as an inspirational role isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatter

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PrefaceThe first edition of the Textbook of Medical Physiologywas written by Arthur C. Guyton almost 55 years ago.Unlike most major medical textbooks, which often have20 or more authors, the first eight editions of the Textbookof Medical Physiology were written entirely by Dr. Guyton,with each new edition arriving on schedule for nearly 40years. The Textbook of Medical Physiology, first publishedin 1956, quickly became the best-selling medical physiology textbook in the world. Dr. Guyton had a gift forcommunicating complex ideas in a clear and interestingmanner that made studying physiology fun. He wrote thebook to help students learn physiology, not to impress hisprofessional colleagues.I worked closely with Dr. Guyton for almost 30 yearsand had the privilege of writing parts of the 9th and 10theditions. After Dr. Guyton’s tragic death in an automobileaccident in 2003, I assumed responsibility for completingthe 11th edition.For the 12th edition of the Textbook of MedicalPhysiology, I have the same goal as for previous editions—to explain, in language easily understood by students, howthe different cells, tissues, and organs of the human bodywork together to maintain life.This task has been challenging and fun because ourrapidly increasing knowledge of physiology continues tounravel new mysteries of body functions. Advances inmolecular and cellular physiology have made it possible to explain many physiology principles in the terminology of molecular and physical sciences rather thanin merely a series of separate and unexplained biologicalphenomena.The Textbook of Medical Physiology, however, is nota reference book that attempts to provide a compendium of the most recent advances in physiology. This isa book that continues the tradition of being written forstudents. It focuses on the basic principles of physiology needed to begin a career in the health care professions, such as medicine, dentistry and nursing, as wellas graduate studies in the biological and health sciences.It should also be useful to physicians and health careprofessionals who wish to review the basic principlesneeded for understanding the pathophysiology ofhuman disease.I have attempted to maintain the same unified organization of the text that has been useful to students inthe past and to ensure that the book is comprehensiveenough that students will continue to use it during their professional careers.My hope is that this textbook conveys the majesty ofthe human body and its many functions and that it stimulates students to study physiology throughout theircareers. Physiology is the link between the basic sciencesand medicine. The great beauty of physiology is that itintegrates the individual functions of all the body’s different cells, tissues, and organs into a functional whole, thehuman body. Indeed, the human body is much more thanthe 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 theseparate organs and systems coordinated to maintainproper function of the entire body? Fortunately, our bodies are endowed with a vast network of feedback controls that achieve the necessary balances without whichwe would be unable to live. Physiologists call this highlevel of internal bodily control homeostasis. In diseasestates, functional balances are often seriously disturbedand homeostasis is impaired. 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 theeffectiveness and beauty of the body’s homeostasis mechanisms as well as to present their abnormal functions indisease.Another objective is to be as accurate as possible.Suggestions and critiques from many students, physiologists, and clinicians throughout the world have beensought and then used to check factual accuracy as well asbalance in the text. Even so, because of the likelihood oferror in sorting through many thousands of bits of information, I wish to issue a further request to all readers tosend along notations of error or inaccuracy. Physiologistsunderstand the importance of feedback for proper function of the human body; so, too, is feedback important forprogressive improvement of a textbook of physiology. Tothe many persons who have already helped, I express sincere p?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmattervii

PrefaceA brief explanation is needed about several features ofthe 12th edition. Although many of the chapters have beenrevised to include new principles of physiology, the textlength has been closely monitored to limit the book sizeso that it can be used effectively in physiology courses formedical students and health care professionals. Many of thefigures have also been redrawn and are in full color. New references have been chosen primarily for their presentationof physiologic principles, for the quality of their own references, and for their easy accessibility. The selected biblio graphy at the end of the chapters lists papers mainly fromrecently published scientific journals that can be freelyaccessed from the PubMed internet site at http://www.ncbi.nlm.nih.gov/sites/entrez/. Use of these references, aswell as cross-references from them, can give the studentalmost complete coverage of the entire field of physiology.The effort to be as concise as possible has, unfortunately,necessitated a more simplified and dogmatic presentationof many physiologic principles than I normally would havedesired. However, the bibliography can be used to learnmore about the controversies and unanswered questionsthat remain in understanding the complex functions of thehuman body in health and disease.Another feature is that the print is set in two sizes. Thematerial in large print constitutes the fundamental physiologic information that students will require in virtuallyall of their medical activities and studies.The material in small print is of several different kinds:first, anatomic, chemical, and other information that isneeded for immediate discussion but that most studentswill learn in more detail in other courses; second, physiologic information of special importance to certain fieldsof clinical medicine; and, third, information that will be ofvalue to those students who may wish to study particularphysiologic mechanisms more deeply.I wish to express sincere thanks to many persons whohave helped to prepare this book, including my colleaguesin the Department of Physiology and Biophysics at theUniversity of Mississippi Medical Center who providedvaluable suggestions. The members of our faculty and abrief description of the research and educational activities of the department can be found at the web site: http://physiology.umc.edu/. I am also grateful to StephanieLucas and Courtney Horton Graham for their excellentsecretarial services, to Michael Schenk and Walter (Kyle)Cunningham for their expert artwork, and to WilliamSchmitt, Rebecca Gruliow, Frank Morales, and the entireElsevier Saunders team for continued editorial and production excellence.Finally, I owe an enormous debt to Arthur Guytonfor the great privilege of contributing to the Textbook ofMedical Physiology, for an exciting career in physiology,for his friendship, and for the inspiration that he providedto all who knew him.John E. sbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatterviii

ContentsUNIT IApoptosis—Programmed Cell DeathCancerIntroduction to Physiology: The Cell andGeneral Physiology4040UNIT IICHAPTER 1Functional Organization of the Human Bodyand Control of the “Internal Environment” Cells as the Living Units of the BodyExtracellular Fluid—The “InternalEnvironment”“Homeostatic” Mechanisms of the MajorFunctional SystemsControl Systems of the BodySummary—Automaticity of the BodyCHAPTER 2The Cell and Its Functions Organization of the CellPhysical Structure of the CellComparison of the Animal Cell withPrecellular Forms of LifeFunctional Systems of the CellLocomotion of CellsCHAPTER 3Genetic Control of Protein Synthesis, CellFunction, and Cell Reproduction Genes in the Cell NucleusThe DNA Code in the Cell Nucleus IsTransferred to an RNA Code in the CellCytoplasm—The Process of TranscriptionSynthesis of Other Substances in the CellControl of Gene Function and BiochemicalActivity in CellsThe DNA-Genetic System Also Controls CellReproductionCell DifferentiationMembrane Physiology, Nerve, and Muscle33346911111217182327273035353739CHAPTER 4Transport of Substances Through CellMembranes The Lipid Barrier of the Cell Membrane,and Cell Membrane Transport ProteinsDiffusion“Active Transport” of Substances ThroughMembranesCHAPTER 5Membrane Potentials and Action Potentials Basic Physics of Membrane PotentialsMeasuring the Membrane PotentialResting Membrane Potential of NervesNerve Action PotentialRoles of Other Ions During the ActionPotentialPropagation of the Action PotentialRe-establishing Sodium and PotassiumIonic Gradients After Action Potentials AreCompleted—Importance of EnergyMetabolismPlateau in Some Action PotentialsRhythmicity of Some Excitable Tissues—Repetitive DischargeSpecial Characteristics of Signal Transmissionin Nerve TrunksExcitation—The Process of Eliciting theAction PotentialRecording Membrane Potentials andAction /www.us.elsevierhealth.com/product.jsp?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatterix

ContentsCHAPTER 6Contraction of Skeletal Muscle Physiologic Anatomy of Skeletal MuscleGeneral Mechanism of Muscle ContractionMolecular Mechanism of Muscle ContractionEnergetics of Muscle ContractionCharacteristics of Whole MuscleContractionCHAPTER 7Excitation of Skeletal Muscle:Neuromuscular Transmission andExcitation-Contraction Coupling Transmission of Impulses from Nerve Endingsto Skeletal Muscle Fibers: The NeuromuscularJunctionMolecular Biology of Acetylcholine Formationand ReleaseDrugs That Enhance or Block Transmissionat the Neuromuscular JunctionMyasthenia Gravis Causes Muscle ParalysisMuscle Action PotentialExcitation-Contraction CouplingCHAPTER 1171717374787983838686868788CHAPTER 8Excitation and Contraction of Smooth Muscle 91Contraction of Smooth Muscle91Nervous and Hormonal Control of SmoothMuscle Contraction94UNIT IIIThe HeartCHAPTER 9Cardiac Muscle; The Heart as a Pump andFunction of the Heart ValvesPhysiology of Cardiac MuscleCardiac CycleRelationship of the Heart Sounds to HeartPumpingWork Output of the HeartChemical Energy Required for Cardiac Contraction:Oxygen Utilization by the HeartRegulation of Heart PumpingCHAPTER 10Rhythmical Excitation of the HeartSpecialized Excitatory and Conductive Systemof the HeartControl of Excitation and Conduction in theHeart101101104107107109110115115118The Normal ElectrocardiogramCharacteristics of the NormalElectrocardiogramMethods for Recording ElectrocardiogramsFlow of Current Around the Heartduring the Cardiac CycleElectrocardiographic LeadsCHAPTER 12Electrocardiographic Interpretation ofCardiac Muscle and Coronary Blood FlowAbnormalities: Vectorial Analysis Principles of Vectorial Analysis ofElectrocardiogramsVectorial Analysis of the NormalElectrocardiogramMean Electrical Axis of the VentricularQRS—and Its SignificanceConditions That Cause Abnormal Voltagesof the QRS ComplexProlonged and Bizarre Patterns of the QRSComplexCurrent of InjuryAbnormalities in the T WaveCHAPTER 13Cardiac Arrhythmias and TheirElectrocardiographic Interpretation Abnormal Sinus RhythmsAbnormal Rhythms That Result from Blockof Heart Signals Within the IntracardiacConduction PathwaysPremature ContractionsParoxysmal TachycardiaVentricular FibrillationAtrial FibrillationAtrial FlutterCardiac 3144146148149151152153UNIT IVThe CirculationCHAPTER 14Overview of the Circulation; Biophysics ofPressure, Flow, and Resistance Physical Characteristics of the CirculationBasic Principles of Circulatory FunctionInterrelationships of Pressure, Flow, lth.com/product.jsp?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatterx

ContentsCHAPTER 15Vascular Distensibility and Functions of theArterial and Venous Systems Vascular DistensibilityArterial Pressure PulsationsVeins and Their FunctionsCHAPTER 16The Microcirculation and LymphaticSystem: Capillary Fluid Exchange,Interstitial Fluid, and Lymph Flow Structure of the Microcirculationand Capillary SystemFlow of Blood in the Capillaries—VasomotionExchange of Water, Nutrients, and OtherSubstances Between the Blood andInterstitial FluidInterstitium and Interstitial FluidFluid Filtration Across Capillaries IsDetermined by Hydrostatic and ColloidOsmotic Pressures, as Well as CapillaryFiltration CoefficientLymphatic SystemCHAPTER 17Local and Humoral Control of TissueBlood Flow Local Control of Blood Flow in Response toTissue NeedsMechanisms of Blood Flow ControlHumoral Control of the CirculationCHAPTER 18Nervous Regulation of the Circulation,and Rapid Control of Arterial Pressure Nervous Regulation of the CirculationRole of the Nervous System in RapidControl of Arterial PressureSpecial Features of Nervous Controlof Arterial PressureCHAPTER 19Role of the Kidneys in Long-Term Control ofArterial Pressure and in Hypertension: TheIntegrated System for Arterial PressureRegulation Renal–Body Fluid System for ArterialPressure ControlThe Renin-Angiotensin System: Its Rolein Arterial Pressure ControlSummary of the Integrated, MultifacetedSystem for Arterial Pressure 91199201201204209213213220226CHAPTER 20Cardiac Output, Venous Return,and Their Regulation Normal Values for Cardiac Output at Restand During ActivityControl of Cardiac Output by VenousReturn—Role of the Frank-Starling Mechanismof the HeartPathologically High or Low Cardiac OutputsMethods for Measuring CardiacOutputCHAPTER 21Muscle Blood Flow and Cardiac OutputDuring Exercise; the Coronary Circulationand Ischemic Heart Disease Blood Flow Regulation in Skeletal Muscleat Rest and During ExerciseCoronary CirculationCHAPTER 22Cardiac Failure Circulatory Dynamics in Cardiac FailureUnilateral Left Heart FailureLow-Output Cardiac Failure—Cardiogenic ShockEdema in Patients with Cardiac FailureCardiac ReserveCHAPTER 23Heart Valves and Heart Sounds;Valvular and Congenital HeartDefects Heart SoundsAbnormal Circulatory Dynamics in ValvularHeart DiseaseAbnormal Circulatory Dynamicsin Congenital Heart DefectsUse of Extracorporeal Circulation DuringCardiac SurgeryHypertrophy of the Heart in Valvularand Congenital Heart DiseaseCHAPTER 24Circulatory Shock and Its Treatment Physiologic Causes of ShockShock Caused by Hypovolemia—Hemorrhagic ShockNeurogenic Shock—Increased VascularCapacityAnaphylactic Shock and Histamine ShockSeptic evierhealth.com/product.jsp?isbn 9781416045748&elsca1 doodys&elsca2 PDF&elsca3 Hall9781416045748&elsca4 frontmatterxi

ContentsPhysiology of Treatment in ShockCirculatory Arrest280281UNIT VThe Body Fluids and KidneysCHAPTER 25The Body Fluid Compartments: Extracellularand Intracellular Fluids; Edema Fluid Intake and Output Are BalancedDuring Steady-State ConditionsBody Fluid CompartmentsExtracellular Fluid CompartmentBlood VolumeConstituents of Extracellular and IntracellularFluidsMeasurement of Fluid Volumes in the DifferentBody Fluid Compartments—the IndicatorDilution PrincipleDetermination of Volumes of Specific BodyFluid CompartmentsR

the 11th edition. For the 12th edition of the Textbook of Medical Physiology, I have the same goal as for 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 fun because our

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