LWBK942-FM.qxd 6/25/11 8:45 AM Page x ACSM’s Advanced Exercise Physiology Second Edition FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd ii 77//88//1111 1122::0022::1188 AAMM Acquisitions Editor: Emily Lupash Product Manager: Andrea Klingler Marketing Manager: Christen Murphy Designer: Doug Smock Compositor: Absolute Service, Inc. Second Edition Copyright © 2012 American College of Sports Medicine 351 West Camden Street Two Commerce Square Baltimore, MD 21201 2001 Market Street Philadelphia, PA 19103 Printed in China All rights reserved. This book is protected by copyright. No part of this book may be reproduced or transmitted in any form or by any means, including as photocopies or scanned-in or other electronic copies, or utilized by any information storage and retrieval system without written permission from the copyright owner, except for brief quotations embodied in critical articles and reviews. Materials appearing in this book prepared by individuals as part of their offi cial duties as U.S. government employees are not covered by the above-mentioned copyright. To request permission, please contact Lippincott Williams & Wilkins at Two Commerce Square, 2001 Market Street, Philadelphia, PA 19103, via email at [email protected], or via website at lww.com (products and services). 9 8 7 6 5 4 3 2 1 Library of Congress Cataloging-in-Publication Data ACSM’s advanced exercise physiology. — 2nd ed. p. ; cm. Advanced exercise physiology / editors, Peter A. Farrell, Michael Joyner, Vincent Caiozzo. Includes bibliographical references and index. ISBN 978-0-7817-9780-1 (alk. paper) 1. Exercise—Physiological aspects. I. Farrell, Peter A. II. Joyner, Michael J. III. Caiozzo, Vincent. IV. American College of Sports Medicine. V. Title: Advanced exercise physiology / editors, Peter A. Farrell, Michael Joyner, Vincent Caiozzo. [DNLM: 1. Exercise—physiology. 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Lippincott Williams & Wilkins cus- tomer service representatives are available from 8:30 AM to 6:00 PM, EST. FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd iiii 77//88//1111 1122::0022::1199 AAMM EDITORS Peter A. Farrell, PhD, FACSM Department of Kinesiology East Carolina University Greenville, NC Michael J. Joyner, MD, FACSM Department of Anesthesiology Mayo Clinic Rochester, MN Vincent J. Caiozzo, PhD, FACSM Departments of Orthopaedic Surgery and Physiology and Biophysics College of Medicine University of California Irvine, CA iii FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd iiiiii 77//88//1111 1122::0022::1199 AAMM FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd iivv 77//88//1111 1122::0022::1199 AAMM PREFACE Exercise physiology is similar to other disciplines that The chapters provide an in-depth summary of how each have matured not only through an exponential expan- system functions and is regulated during the physiologi- sion of the literature but also in the need for researchers cal stress of exercise. Our goal in this second edition was to specialize in particular systems or functions. This need to take advantage of the fact that all of the authors are the to specialize, however, poses a fundamental challenge in scientists that a graduate student should seek out when the education of the next generation of exercise physiolo- they have a need for detailed information and understand- gists: there remains a need for an advanced understanding ing in the various areas. We have anticipated those needs of all systems because of the simple fact that each of the and have reorganized the chapters so that the major points body systems interacts with and, in many cases, regulates have been highlighted and presented in terms that a typi- other systems. It seems quite unlikely that even talented cal graduate/upper-level undergraduate student in exercise younger students could read a journal article about specif- physiology could comprehend. ics of frequency modulation of motor units and come away with a satisfying understanding of data being presented in Organization a primary literature paper. The American College of Sports Medicine’s (ACSM’s) Advanced Exercise Physiology, Second Edition, bridges the gap Students normally seek graduate programs whose fac- between that basic understanding of physiology during ulty has expertise in specifi c areas of exercise physiology. exercise (knowledge gained in an introductory course on Unfortunately, most graduate programs do not have suf- exercise physiology) and information presented through fi cient breadth of faculty expertise to teach all the areas the primary literature in high-quality journals. The chap- included in this textbook. Information in all chapters is ters are written by authorities in most of the major areas at a level that, once understood, will assure that students of investigation in exercise physiology, and the content in possess up-to-date, coherent, and accurate information at this edition has been updated to include the latest facts an advanced level. and concepts available to authorities in the fi eld. Each This second edition of Advanced Exercise Physiology has chapter presents a detailed summary of facts and solidi- been reorganized so that each chapter is presented in a simi- fying concepts necessary for students to embrace before lar manner, which makes the information fl ow more com- they can understand the specifi cs in the primary literature. prehensibly for students. Though the chapters continue to v FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd vv 77//88//1111 1122::0022::1199 AAMM vi Preface center on specifi c topics, the second edition cross-references The fi nal section of the book centers on how environ- points made that involve concepts/facts pertinent to more mental factors infl uence many of the regulatory mecha- than one chapter. This approach emphasizes the fundamen- nisms discussed in the previous chapters. Separate chapters tal appreciation that no physiological system functions in discuss heat, cold, hypoxia, hyperbaria, and exercise in isolation especially during exercise. space. Chapter 27 is quite futuristic in nature; it covers ACSM’s Advanced Exercise Physiology, Second Edition, exercise genomics and proteomics from the perspective of begins with an introductory chapter on the history of exer- how modern techniques can be used to better understand cise physiology that provides excellent insights into how how exercise alters gene expression and more. the science of exercise physiology developed through a combination of medicine and, at fi rst, preparation for war Features or fi ghting and then physical performance in the Olympics and international competitions. Subsequent chapters dis- cuss specifi c systems such as nervous system control of New additions to the chapters include Key Points, which movement, muscle architecture, force generation, and, highlight the most important concepts or facts, and section when these processes fail, the fatigue process. Perhaps the Summaries, which provide brief synopses of key sections. most extensive single chapter is Chapter 8, “The Respi- We also include Test Yourself questions, which appear ratory System,” which presents a detailed description of throughout the chapters and will require students to use respiratory function and the known causes for increased critical thinking skills to discuss hypotheses and how the ventilation during exercise, as well as other aspects of lung information they have learned relates to practical appli- function. Extensive coverage of the cardiovascular system is cations. We are hopeful these questions also will help to provided in three chapters (C hapters 9–11) that discuss the stimulate class or group discussions. structure of the cardiovascular system, heart function, and circulatory control at the level of the microcirculation. Additional Resources An extensive section on metabolism begins with a ACSM’s Advanced Exercise Physiology, Second Edition, includes review of how the gastrointestinal system functions to additional resources for both instructors and students that deliver nutrients before, during, and after exercise and are available on the book’s companion website at http:// its role in maintaining acid–base, electrolyte, and water thepoint.lww.com/ACSMAdvExPhys2e. balance during exercise (Chapter 12). Because of the im- portance of providing fuels during exercise, six chapters Instructors (Chapters 13–18) are devoted to carbohydrate, fat, and pro- tein metabolism during exercise and in response to regu- • Image bank lar exercise training. Chapter 19, “The Endocrine System: • PowerPoint lecture outlines Integrated Infl uences on Metabolism, Growth, and Repro- • WebCT/Blackboard/Angel LMS cartridge duction,” provides coverage of many hormonal regulators that control several systems described in other chapters, Students as well as exercise metabolism. An additional strength • Full text online of this chapter is the coverage of endocrine changes that • Interactive quiz bank alter reproductive function, with a primary, but not ex- • Animations clusive, emphasis on women. The chapter on immunol- ogy provides the latest information on how the immune In addition, purchasers of the text can access the system responds to exercise and equally important how searchable full text through the book’s website. See the the immune system interacts with exercise metabolism. front inside cover of this text for more details and include This chapter also reviews literature supporting the con- the passcode to gain access to the website. cept of muscle being an endocrine organ. This section of the book culminates with a discussion on body fl uid con- Peter Farrell trol mechanisms during exercise and includes the renal Mike Joyner system (Chapters 20–22). Vince Caiozzo FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd vvii 77//88//1111 1122::0022::1199 AAMM CONTRIBUTORS Charles Tipton, PhD Gregory R. Wohl, PhD, PEng Department of Physiology Department of Mechanical Engineering University of Arizona McMaster University Tucson, AZ Hamilton, Ontario, Canada Chapter 1 Chapter 3 V. Reggie Edgerton, PhD Jeremy M. LaMothe, MD, PhD Department of Integrative Biology and Physiology Department of Surgery Department of Neurobiology University of Calgary Brain Research Institute Calgary, Alberta, Canada University of California, Los Angeles Chapter 3 Los Angeles, CA Chapter 2 Vincent J. Caiozzo, PhD Departments of Orthopaedic Surgery and Physiology Roland R. Roy, PhD, FACSM and Biophysics Department of Integrative Biology and Physiology College of Medicine Brain Research Institute University of California University of California, Los Angeles Irvine, CA Los Angeles, CA Chapter 4 Chapter 2 Richard Tsika, PhD Ronald F. Zernicke, PhD, DSc, FACSM Department of Biomedical Sciences School of Kinesiology University of Missouri Departments of Orthopaedic Surgery and Biomedical Columbia, MO Engineering Chapter 5 University of Michigan Ann Arbor, MI Chapter 3 vii FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd vviiii 77//88//1111 1122::0022::1199 AAMM viii Contributors Robert H. Fitts, PhD David A. Brown, PhD Department of Biological Sciences Department of Physiology Marquette University East Carolina University Milwaukee, WI Greenville, NC Chapter 6 Chapter 10 Douglas R. Seals, PhD Steven S. Segal, PhD, FACSM Department of Integrative Physiology Department of Medical Pharmacology and Physiology University of Colorado at Boulder University of Missouri–Columbia Boulder, CO Columbia, MO Chapter 7 Chapter 11 Lee M. Romer, PhD, FACSM Shawn E. Bearden, PhD, FAHA Reader (Human and Applied Physiology) Department of Biological Sciences Centre for Sports Medicine and Human Performance Biomedical Research Institute Brunel University Idaho State University Uxbridge Middlesex, UK Pocatello, ID Chapter 8 Chapter 11 A. William Sheel, PhD G. Patrick Lambert, PhD School of Human Kinetics Department of Exercise Science University of British Columbia Creighton University Vancouver, British Columbia, Canada Omaha, NE Chapter 8 Chapter 12 Craig A. Harms, PhD, FACSM Xiaocai Shi, PhD, FACSM Department of Kinesiology Gatorade Sports Science Institute Kansas State University Barrington, IL Manhattan, KS Chapter 12 Chapter 8 Donal S. O’Leary, PhD Robert Murray, PhD, FACSM Department of Physiology Sports Science Insights, LLC Wayne State University Crystal Lake, IL School of Medicine Chapter 12 Detroit, MI Chapter 9 Ronald A. Meyer, PhD Departments of Physiology and Radiology Patrick J. Mueller, PhD Michigan State University Department of Physiology East Lansing, MI Wayne State University School of Medicine Chapter 13 Detroit, MI Chapter 9 Robert W. Wiseman, PhD Departments of Physiology and Radiology Javier A. Sala-Mercado, MD, PhD Michigan State University Department of Physiology East Lansing, MI Cardiovascular Research Institute Chapter 13 Wayne State University, School of Medicine Detroit, MI Mark Hargreaves, PhD Chapter 9 Department of Physiology University of Melbourne Russell L. Moore, PhD Melbourne, Australia Department of Integrative Physiology Chapter 14 University of Colorado at Boulder Boulder, CO Chapter 10 FFaarrrreellll__2288336666__FFMM__PPrriinntteerr__FFiillee..iinndddd vviiiiii 77//88//1111 1122::0022::1199 AAMM