Medical Physiology
WALTER F. BORON, MD, PhD Professor David N. and Inez Myers/Antonio Scarpa Chairman Department of Physiology and Biophysics Case Western Reserve University Cleveland, Ohio
EMILE L. BOULPAEP, MD Professor Department of Cellular and Molecular Physiology Yale University School of Medicine New Haven, Connecticut
MEDICAL PHYSIOLOGY, THIRD EDITION INTERNATIONAL EDITION Copyright 2017 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. Details on how to seek permission, further information about the Publisher's permissions policies and our arrangements with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: wwwelsevier.com/permissions. This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted herein).
Notices Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary. Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any information, methods, compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the safety of others, including panties for whom they have a professional responsibility. With respect to any drug or pharmaceutical products identified, 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 practitioners, relying on their own experience and knowledge of their patients, 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 authors, contributors, or editors, assume any liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein. Previous editions copyrighted 2012, 2009, 2005, and 2003. Library of Congress Cataloging-in-Publication Data Names: Boron, Walter E, editor. 1 Boulpaep, Emile L., editor. Title: Medical physiology / [edited by] Walter E Boron, Emile L. Boulpaep. Other titles: Medical physiology (Boron) Description: Edition 3.1 Philadelphia, PA : Elsevier, [2017] 1 Includes bibliographical referentes and Ăndex. Identifiers: LCCN 20160052601 ISBN 9781455743773 (hardcover alk. paper) ISBN 9780323427968 (International ed.) Subjects: MESH: Physiological Phenomena 1 Cell Physiological Phenomena Classification: LCC QP34.5 1 NLM QT 104 1 DDC 612—dc23 LC record available at http://lccn.loc.gov/2016005260
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CONTENTS SECTION 1 INTRODUCTION 1 Foundations of Physiology, 2 Emile L. Boulpaep and Walter F. Boron
SECTION IV THE CARDIOVASCULAR SYSTEM 17 Organization of the Cardiovascular System, 410 Emile L Boulpaep
18 Blood, 429 Emile L. Boulpaep
SECTION II
19 Arteries and Veins, 447
PHYSIOLOGY OF CELLS AND MOLECULES
20 The Microcirculation, 461
2 Functional Organization of the Cell, 8 Michael J. Caplan
3 Signal Transduction, 47 Lloyd Cantley
4 Regulation of Gene Expression, 73 Peter lgarashi
5 Transport of Solutes and Water, 102 Peter S. Aronson, Walter F. Boron, and Emile L. Boulpaep
6 Electrophysiology of the Cell Membrane, 141
Edward G. Moczydlowski 7 Electrical Excitability and Action Potentials, 173 Edward G. Moczydlowski
8 Synaptic Transmission and the Neuromuscular Junction, 204
Emile L Boulpaep Emile L Boulpaep
21 Cardiac Electrophysiology and the Electrocardiogram, 483 W. Jonathan Lederer
22 The Heart as a Pump, 507 Emile L. Boulpaep
23 Regulation of Arterial Pressure and Cardiac O utp ut, 533 Emile L. Boulpaep
24 Special Circulations, 556 Steven S. Segal
25 Integrated Control of the Cardiovascular System, 572
Emile L. Boulpaep
Edward G. Moczydlowski 9 Cellular Physiology of Skeletal, Cardiac, and Smooth
Muscle, 228 Edward G. Moczydlowski
SECTION V THE RESPIRATORY SYSTEM 26 Organization of the Respiratory System, 590
SECTION 111 THE NERVOUS SYSTEM 10Organization of the Nervous System, 254 Bruce R. Ransom 11 The Neuronal Microenvironment, 275 Bruce R. Ransom 12 Physiology of Neurons, 295 Barry W. Connors 13 Synaptic Transmission in the Nervous System, 307 Barry W. Connors
14 The Autonomic Nervous System, 334 George B. Richerson 15 Sensory Transduction, 353
Walter F. Boron
27 Mechanics of Ventilation, 606 Walter F. Boron
28 Acid-Base Physiology, 628 Walter F. Boron
29 Transport of Oxygen and Carbon Dioxide in the Blood, 647 Walter F. Boron
30 Gas Exchange in the Lungs, 660 Walter F. Boron
31 Ventilation and Perfusion of the Lungs, 675 Walter F. Boron
32 Control of Ventilation, 700 George B. Richerson and Walter F. Boron
Barry W. Connors 16 Circuits of the Central Nervous System, 390 Barry W. Connors
xi
SECTION VI THE URINARY SYSTEM 33 Organization of the Urinary System, 722 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson
34 Glomerular Filtration and Renal Blood Flow, 739 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson
35 Transport of Sodium and Chloride, 754 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson
36 Transport of Urea, Glucose, Phosphate, Calcium, 37
Magnesium, and Organic Solutes, 770 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson Transport of Potassium, 792 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson
38 Urine Concentration and Dilution, 806 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson
39 Transport of Acids and Bases, 821 40
Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson Integration of Salt and Water Balance, 836 Gerhard Giebisch, Erich E. Windhager, and Peter S. Aronson
SECTION VII THE GASTROINTESTINAL SYSTEM
48 Endocrine Regulation of Growth and Body Mass, 990 Eugene J. Barrett
49 The Thyroid Gland, 1006 Eugene J. Barrett
50 The Adrenal Gland, 1018 Eugene J. Barrett
51 The Endocrine Pancreas, 1035 Eugene J. Barrett
52 The Parathyroid Glands and Vitamin D, 1054 Eugene J. Barrett and Paula Q. Barrett
SECTION
IX
THE REPRODUCTIVE SYSTEM 53 Sexual Differentiation, 1072 Sam Mesiano and Ervin E. Jones
54 The Male Reproductive System, 1092 Sam Mesiano and Ervin E. Jones
55 The Female Reproductive System, 1108 Sam Mesiano and Ervin E. Jones 56 Fertilization, Pregnancy, and Lactation, 1129 Sam Mesiano and Ervin E. Jones 57 Fetal and Neonatal Physiology, 1151 George Lister and Ervin E. Jones
41 Organization of the Gastrointestinal System, 852 Henry J. Binder 42 Gastric Function, 863 Henry J. Binder 43 Pancreatic and Salivary Glands, 879 Christopher R. Marino and Fred S. Gorelick 44 Intestinal Fluid and Electrolyte Movement, 899 Henry J. Binder 45 Nutrient Digestion and Absorption, 914 Henry J. Binder and Charles M. Mansbach II 46 Hepatobiliary Function, 944 Frederick J. Suchy
SECTION VIII THE ENDOCRINE SYSTEM 47 Organization of Endocrine Control, 974 Eugene J. Barrett
SECTION X PHYSIOLOGY OF EVERYDAY LIFE 58 Metabolism, 1170 Gerald I. Shulman and Kitt Falk Petersen 59 Regulation of Body Temperature, 1193 Shaun F. Morrison 60 Exercise Physiology and Sports Science, 1204 Steven S. Segal 61 Environmental Physiology, 1223 Arthur DuBois 62 The Physiology of Aging, 1235 Edward J. Masoro Index, 1249