Exercise Physiology: Nutrition, Energy, and Human Performance [7. ed] 9780781797818, 0781797810

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EXERCISE PHYSIOLOGY Nutrition, Energy, and Human Performance Seventh Edition

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EXERCISE PHYSIOLOGY Nutrition, Energy, and Human Performance Seventh Edition

William D. McArdle Professor Emeritus, Department of Family, Nutrition, and Exercise Science Queens College of the City University of New York Flushing, New York

Frank I. Katch International Research Scholar Faculty of Health and Sport, Agder University College Kristiansand, Norway Instructor and Board Member Certificate Program in Fitness Instruction UCLA Extension, Los Angeles, California Former Professor and Chair of Exercise Science University of Massachusetts, Amherst, Massachusetts

Victor L. Katch Professor, Department of Movement Science Division of Kinesiology Associate Professor, Pediatrics School of Medicine University of Michigan Ann Arbor, Michigan

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Acquisitions Editor: Emily Lupash Product Manager: Andrea M. Klingler Marketing Manager: Christen Murphy Production Editor: Paula C. Williams Vendor Manager: Kevin Johnson Design Coordinator: Stephen Druding Compositor: Macmillan Publishing Solutions Seventh Edition Copyright © 2010, 2007, 2001, 1996, 1991, 1986, 1981 by Lippincott Williams & Wilkins, a Wolters Kluwer business. 351 West Camden Street Baltimore, MD 21201

530 Walnut Street Philadelphia, PA 19106

Printed in China Unless otherwise indicated, all photographs are Copyright © 2008 by Fitness Technologies, Inc., Frank I. Katch, and Victor L. Katch, 5043 Via Lara Lane, Santa Barbara, CA 93111. This material is protected by copyright. No photograph may be reproduced in any form or by any means without permission from the copyright holders. Appendices B and C Copyright © 2008 by Frank I. Katch, Victor L. Katch, William D. McArdle, and Fitness Technologies, Inc., 5043 Via Lara Lane, Santa Barbara, CA 93111. No part of this appendix may be reproduced in any manner or by any means without written permission from the copyright holders. 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 official duties as U.S. government employees are not covered by the above-mentioned copyright. To request permission, please contact Lippincott Williams & Wilkins at 530 Walnut Street, Philadelphia, PA 19106, 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 McArdle, William D. Exercise physiology : nutrition, energy, and human performance / William D. McArdle, Frank I. Katch, Victor L. Katch. — 7th ed. p. ; cm. Includes bibliographical references and index. ISBN 978-0-7817-9781-8 (alk. paper) 1. Exercise—Physiological aspects. I. Katch, Frank I. II. Katch, Victor L. III. Title. [DNLM: 1. Exercise—physiology. 2. Nutritional Physiological Phenomena. 3. Sports Medicine. QT 260 M478e 2010] QP301.M375 2010 612'.044—dc22 2009029341 DISCLAIMER Care has been taken to confirm the accuracy of the information present and to describe generally accepted practices. However, the authors, editors, and publisher are not responsible for errors or omissions or for any consequences from application of the information in this book and make no warranty, expressed or implied, with respect to the currency, completeness, or accuracy of the contents of the publication. Application of this information in a particular situation remains the professional responsibility of the practitioner; the clinical treatments described and recommended may not be considered absolute and universal recommendations. The authors, editors, and publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accordance with the current recommendations and practice at the time of publication. However, in view of ongoing research, changes in government regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any change in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new or infrequently employed drug. Some drugs and medical devices presented in this publication have Food and Drug Administration (FDA) clearance for limited use in restricted research settings. It is the responsibility of the health care provider to ascertain the FDA status of each drug or device planned for use in their clinical practice. To purchase additional copies of this book, call our customer service department at (800) 638-3030 or fax orders to (301) 223-2320. International customers should call (301) 223-2300. Visit Lippincott Williams & Wilkins on the Internet: http://www.lww.com. Lippincott Williams & Wilkins customer service representatives are available from 8:30 am to 6:00 pm, EST.

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To my wife Kathleen, my best friend and biggest supporter, and to the rest of the “A team”: my children, Theresa, Amy, Kevin, and Jennifer; their spouses, Christian, Jeff, Nicole, and Andy; and my grandchildren, Liam, Aidan, Quinn, Dylan, Kelly Rose, Owen, Henry, Kathleen (Kate), Grace, Elizabeth, Claire, and Elise. Bill McArdle

To my wife and life partner, Kerry, for four decades of love and support; to my two sons, David and Kevin, for achieving the honorable in their business and personal lives; to my daughter, Ellen, for current and future years of dedicated service as a pediatrician; and to her new husband, Sean. Frank Katch

To those most important in my life: Heather, Erika, Leslie, and the J-man. I’m a lucky man. Victor Katch

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PREFACE

As in the publication of the first edition of Exercise Physiology: Nutrition, Energy, and Human Performance in 1981, this seventh edition reflects our continued commitment to integrate the concepts and science of the different disciplines that contribute to a more comprehensive understanding and appreciation of modern-day exercise physiology. Consistent with previous editions, we believe firmly that the content domain of exercise physiology must be predicated on the logical and natural flow of basic knowledge concerning nutrient-energy intake, nutrient-energy metabolism, and systems physiology as related to submaximal and maximal human movement. Current concepts of energy balance intimately link the role of physical inactivity to the “creeping obesity” epidemic that continues to afflict the world’s population. We are encouraged that the medical establishment and governmental agencies have finally acknowledged regular physical activity as an important weapon in the armamentarium for prevention and rehabilitation of diverse disease states, including obesity. We are gratified with the small part we have played in the education of more than 350,000 undergraduate and graduate students who have used this text since the publication of the first edition in 1981. A source of great pride for us is that some of the first students enrolled in our own classes that used this text have gone on to earn advanced degrees in the same or similar fields. This tradition of textbook adoption has now been passed down to their students, many of whom comprise the next generation of aspiring teachers, exercise specialists, and researchers. In fact, one of us (VK) has had the opportunity on three different occasions to teach students whose parents were former students. We are forever grateful to our former teachers and mentors for igniting a spark that has not diminished. We hope you will become as excited as we first were (and continue to be) in the science of exercise physiology and human performance.

ORGANIZATION This seventh edition maintains the same seven-section structure as previous editions, including an introductory section about the origins of exercise physiology and a concluding “On the Horizons” section that deals with a maturing effort in exercise physiology to incorporate molecular biology to human performance and the many interrelated aspects of health and disease.

FEATURES Many features throughout the text are included to engage the student and facilitate learning. These include the following:

Introduction: A View of the Past. The text’s introduction, “Exercise Physiology: Roots and Historical Perspectives,” reflects our interest and respect for the earliest underpinnings of the field, and the direct and indirect contributions of the men and women physicians/scientists who contributed to the field. In a Practical Sense. This element in every chapter highlights practical applications that include: • Lowering high blood pressure with dietary intervention: the DASH diet • Leveraging nutrition to prevent chronic athletic fatigue • Predicting V·O2max during pregnancy from submaximal exercise heart rate and oxygen consumption • Predicting energy expenditure during treadmill walking and running • Determining anaerobic power and capacity: the Wingate cycle ergometer test • Predicting pulmonary function variables in males and females • Measuring lactate threshold • Blood pressure measurement, classifications, and recommended follow-up • Placing electrodes of bipolar and 12-lead ECG recordings • Diabetes, hypoglycemia, and exercise • Protecting the lower back • Assessing heat quality of the environment: how hot is too hot? Focus on Research. Each chapter’s Focus on Research features a key research article from a renowned scientist. These well-designed studies illustrate how “theory comes to life” through research. Integrative Questions. Another element in each chapter, “Integrative Questions,” poses open-ended questions to encourage students to consider complex concepts without a single “correct” answer. Expanded Art Program. The full-color art program continues to be an important feature of the textbook. New figures have been added to enhance the new and updated content. Up-Close and Personal Interviews. The text features nine contemporary scientists whose important research contributions and visionary leadership continue the tradition of the scientists of prior generations—Steven Blair, Frank Booth, Claude Bouchard, David Costill, Barbara Drinkwater, John Holloszy, Loring Rowell, Bengt Saltin, and Charles Tipton.

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These individuals clearly merit recognition, not only for expanding knowledge through their many scientific contributions, but also for elucidating mechanisms that underlie responses and adaptations to exercise and health enhancement. Each person has been placed within a section linked to his or her main scholarship interests, yet all of them span one or more sections in terms of scientific contributions. Appendix E, which is available online at http://thepoint.lww. com/mkk7e, lists individual honors and awards for each of these distinguished and meritorious scientist–researchers. The intimate insights from the “superstars” should inspire current exercise physiology students to actualize their potential, whether through accomplishments in graduate school, teaching, research, or numerous other exciting opportunities to achieve excellence. References and Appendices Available Online. All references and appendices are available online at http://thepoint. lww.com/mkk7e. Appendices feature valuable information about nutritive values, energy expenditures, metabolic computations in open-circuit spirometry, and more.

NEW TO THE SEVENTH EDITION The flow of information in this edition remains similar to prior editions. Where applicable, figures, tables, and Web sites have been updated and/or expanded to include the most relevant current information, including new tabular material and illustrations to clarify important concepts and information. Significant Additions and Modifications to the Text • Section 1 summarizes the current (2009) energy, nutrient, and fluid recommendations from the American Dietetic Association, Dietitians of Canada, and the American College of Sports Medicine for active adults and competitive athletes. It also provides an expanded discussion of the efficacy and health benefits of routinely consuming vitamin and mineral supplements versus obtaining these micronutrients in the foods of a well-balanced diet. • In Section 2, we have included the latest information regarding the energy yield from the catabolism of the different macronutrients and we have rewritten several of the sections to increase clarity and specificity. • We have expanded our discussion of the increasing incidence of hypertension with age in Section 3, along with lifestyle choices to lower blood pressure. • In Section 4, we present a research-based alternative method to estimate maximal heart rate from chronological age for adults. Additionally, we present the latest information regarding sling exercise training as a means of muscle activation and overload that is gaining in popularity based on new research about how muscles are activated and contribute to improved movement. We have also expanded our review of the effects of carbohydrate–protein supplement timing

and resistance training on muscle fiber hypertrophy, muscular strength, and body composition. • Section 5 includes a discussion of the “secret” First Lady Astronaut Trainees (FLATS) program intended to include highly experienced female aviators for future space missions, and how that program was unceremoniously scuttled because of bureaucratic cronyism at the highest levels of the early space agency. Also included is an explanation of the longterm United States human and robotic program to explore the solar system, starting with a return to the moon to ultimately enable future exploration of Mars and other destinations, including the new manned exploration, the Crew Exploration Vehicle (CEV). • Section 6 presents the 2009 summary statement of the American College of Sports Medicine as to the appropriate physical activity intervention strategies for weight loss and prevention of weight regain for adults. Also included is a discussion of an apparent anomaly in body proportions in champion swimmer Michael Phelps, winner of 8 gold medals in the 2008 Beijing Olympics, related to his use of a controversial swim suit and swimming speed. This section also analyzes body size (BMI) differences among 1124 Division 1 Big Ten Collegiate offensive and defensive linemen and their teams related to team standings in league play. We provide the first presentation on the height and weight, and BMI of professional male tour Professional golfers (n = 33) and Champions Tour players (n = 18) compared to 257 golfers, stratified by proficiency levels based on handicap index, and 300,818 Swedish golfers (203,778 men and 97,040 women) with stratification for age, sex, and socioeconomic status. We also include the latest information about brown adipose tissue in humans, and its role in metabolism and link to health and disease. • Section 7 presents an updated section on coronary heart disease. Our final chapter discusses the new molecular exercise physiology program at the University of Aberdeen in Scotland, where the MSc program, including Diploma and Certificate programs, offers courses in a new subfield in sports science that focuses on genetics and signal transduction related to exercise. Current statistics reveal the impact of molecular biology-related research on different fields of science. For example, searching on the terms muscle and genes increased from 502 in 2001 to over 58,000 at the start of 2009! We also highlight Darwin’s monumental contributions in evolutionary theory. A new feature of this final chapter links to the LWW website and includes (1) readings related to molecular biology and genetics, twins, and human performance; (2) reference to excellent texts that devote hundreds of pages to the intricacies of the molecular biology of gene transcription and protein synthesis; (3) articles from Scientific American that concern molecular biology; (4) useful molecular

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biology Internet sites; (5) microscope technologies germane to molecular biology (light microscope, fluorescence microscope, electron microscope, and positron emission tomography or PET); (6) reprint of Watson and Crick’s one-page classic paper in Nature about their deduction of DNA’s structure, which nearly six decades later unraveled the pieces to the primordial jigsaw puzzle of the Human Genome Project; and (7) a timeline of events about genetics before Mendel, followed by notable events in genetics and molecular biology to 2005. Our current reference list includes the latest research gleaned from national and international journals related to specific topic areas. We hope you profit from and enjoy this continuation of our journey through the ever-expanding and relevant field of exercise physiology.

ANCILLARIES: THE TOTAL TEACHING PACKAGE Exercise Physiology: Nutrition, Energy, and Human Performance, Seventh Edition, includes additional resources for both instructors and students that are available on the book’s companion website at http://thepoint.lww.com/mkk7e. Instructors and students will have access to animations illustrating the most important concepts in human physiology. Both instructors and students will also be able to access the searchable Full Text Online.

ix

Instructors Approved adopting instructors will be given access to the following additional resources: • Brownstone test generator • PowerPoint presentations: one set with lecture outlines; one set with images only • Image bank • WebCT, Blackboard, and Angel-Ready Cartridge Students Students who have purchased Exercise Physiology: Nutrition, Energy, and Human Performance, Seventh Edition, have access to the following additional resources: • • • • •

Online quiz bank with study and test options Animations References Appendices Featured information on microscope technologies, notable events in genetics, Nobel prizes, and outstanding female scientists.

Ancillaries were prepared by the authors as well as by Jeff Woods (University of Illinois at Urbana-Champaign) and Lamia Scherzinger (Indiana University–Purdue University Indianapolis).

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ACKNOWLEDGMENTS

We wish to thank many individuals. First, Dr. Loring Rowell for his constructive comments on the chapters related to pulmonary and cardiovascular dynamics during rest and exercise, particularly the sections related to the possible role of the venous system as an active vasculature. We thank Drs. Victor Convertino and Charles Tipton for insightful comments and suggestions on the microgravity chapter. Stephen Lee (Exercise Physiology Laboratory, Johnson Space Center, Houston) kindly supplied original NASA photos and documents; Mission Specialist Astronaut Dr. Martin Fettman (Colorado State University, Ft. Collins, CO) provided original slides he took during his Skylab 2 Mission; and Dr. Helen Lane (Chief Nutritionist, Johnson Space Center, Houston) provided prepublication documents and resource materials. Dr. Ron White, National Space Biomedical Research Institute, allowed us to use charts he helped create from Human Physiology In Space. Teacher’s Manual. We sincerely appreciate the expertise of Drs. Frank Booth, University of Missouri, Kristin Steumple, Department of Health and Exercise Science at Gettysburg College, and Marvin Balouyt, Washtenaw Community College, Ann Arbor, MI, for their expert opinions and suggestions for improving the chapter on molecular biology. Hypoxico Inc. provided photos of the Hypoxico altitude tent. Mr. John Selby (www.hyperlite.co.uk) kindly provided timely information and photos of the portable, collapsible decompression chamber. Dr. Alex Knight, York University, UK, graciously provided information about molecular biology techniques he has pioneered (in vitro motility assay) and other information and a photograph about myosin, muscle, and single molecules. Yakl Freedman (www.dna2z.com) was supportive in supplying recent information about DNA and molecular biology. Sue Hilt of the American College of Sports Medicine staff headquarters did a superb job of securing the text of the Citation and Honor Awards reproduced in Appendix E. Dr. James A. Freeman, professor of English, University of Massachusetts, unselfishly lent his expertise to make words sing. Dr. Barry Franklin, Beaumont Hospital, Detroit, MI, supplied original information about cardiac rehabilitation. Paul Petrich, Goleta, CA, provided photos of scuba expeditions. The Trustees of Amherst College and Archival Library gave permission to reproduce the photographs and materials of Dr. Hitchcock. Magnus Müller, the University of Giessen, kindly provided the photo of Liebig’s Giessen lab on p. xxxvi. We are grateful to marine artist Ron Scobie, ASMA (www.ronscobie-marineartist.com), for his kind permission to reproduce his rendering of the HMS Beagle. We are collectively indebted to the nine researchers–scholars who took time from their busy schedules to answer our interview questions and provide personal photos. Each of those individuals, in his or her own unique way, inspired the three of us in our careers by their work ethic, scientific excellence, and generosity of time and advice with colleagues and students. Over the years, we have had the good fortune to

come to know these individuals both socially and in the academic arena. We must admit, however, that the interviews provided insights previously unknown to us. We hope you too are as impressed as we are by all they have accomplished and given back to the profession. Frank Katch also wishes to thank Dr. Drinkwater, who served on his MS thesis committee at UC Santa Barbara. He now ’fesses up after 45 years that she provided much needed statistical and grammatical assistance beyond the call of duty with that project! We give a special thanks to that group of former students who earned doctoral degrees in physical education, exercise science, and medicine, and who have gone on to distinguish themselves as teachers, clinicians, and researchers in the related areas of exercise physiology. These include Denise Agin, Stamatis Agiovlasitis, Doug Ballor, Dan Becque, Christos Balabinis, George Brooks, Barbara Campaigne, Ed Chaloupka, Ken Cohen, Edward Coyle, Dan Delio, Julia Chase Delio, Joseph DeGuzman, Chris Dunbar, Patti Freedson, Roger Glaser, Ellen Glickman, Kati Haltiwinger, Everett Harmon, Jay Hoffman, Tibor Hortobagyi, Mitch Kanter, Betsy Keller, Jie Kang, Marliese Kimmerly, George Lesmes, Steve Lichtman, Charles Marks, Karen Nau-White, Laurel Traeger-Mackinnon, Robert Mofatt, Steve Ostrove, James Rimmer, Deborah Rinaldi, Stan Sady, Michelle Segar, Lapros Sidossis, Bob Spina, John Spring, Bill Thorland, Mike Toner, Lorraine Turcotte, John Villanacci, Jonnis Vrabis, Nancy Weiss, Nancy Wessinger, Stephen Westing, Art Weltman, Anthony Wilcox, and Linda Zwiren. We also acknowledge the following Master’s and senior honors students who contributed so much to our research and personal experiences: Pedro Alexander, Christos Balabinis, Margaret Ballantyne, Brandee Black, Michael Carpenter, Steven Christos, Roman Czula, Gwyn Danielson, Toni Denahan, Marty Dicker, Sadie Drumm, Peter Frykman, Scott Glickman, Marion Gurry, Carrie Hauser, Margie King, Peter LaChance, Jean Lett, Maria Likomitrou, Robert Martin, Cathi Moorehead, Susan Novitsky, Joan Perry, Sharon Purdy, Michelle Segar, Debra Spiak, Lorraine Turcotte, Lori Waiter, Stephen Westing, and Howard Zelaznik. Finally, we would like to thank all of the individuals at LWW who helped bring this seventh edition to fruition. We are particularly indebted to Andrea Klingler, our superb product manager, who continually provided support, subtle urging, and overall excellence in helping to bring this seventh edition to fruition. Thank you, Andrea! William D. McArdle Sound Beach, NY Frank I. Katch Santa Barbara, CA Victor L. Katch Ann Arbor, MI

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CONTENTS

A View of the Past: Exercise Physiology— Roots and Historical Perspectives xvii

Phosphorus 65 Magnesium 65 Iron 67 Sodium, Potassium, and Chlorine 71 Minerals and Exercise Performance 72

INTRODUCTION

Interview with Dr. Charles Tipton

PART ONE: EXERCISE PHYSIOLOGY Section 1

Nutrition: The Base for Human Performance 3

Interview with Dr. David L. Costill CHAPTER 1

Part 3 • WATER 75 The Body’s Water Content 75 Water Balance: Intake Versus Output 77 Water Requirement in Exercise 78

1

CHAPTER 3

Carbohydrates, Lipids, and Proteins 7 Part 1 • CARBOHYDRATES 8 Kinds and Sources of Carbohydrates 8 Recommended Intake of Carbohydrates 13 Role of Carbohydrates in the Body 13 Carbohydrate Dynamics in Exercise 16 Part 2 • LIPIDS 20 The Nature of Lipids 20 Kinds and Sources of Lipids 20 Recommended Lipid Intake 26 Role of Lipid in the Body 27 Fat Dynamics During Exercise 28 Part 3 • PROTEINS 31 The Nature of Proteins 31 Kinds of Protein 31 Recommended Protein Intake 33 Role of Protein in the Body 35 Dynamics of Protein Metabolism 37 Nitrogen Balance 37 Protein Dynamics in Exercise and Training 39

CHAPTER 2

Section 2

Part 2 • MINERALS 56 The Nature of Minerals 56 Role of Minerals in the Body 56 Calcium 56 The Female Athlete Triad: Unexpected Problem for Women Who Train Intensely 63

Energy for Physical Activity 107

Interview with Dr. John O. Holloszy CHAPTER 4

Energy Value of Food 111 Measurement of Food Energy 112

CHAPTER 5

Introduction to Energy Transfer 118 Energy—The Capacity for Work 119 Interconversions of Energy 121 Biologic Work in Humans 123 Factors that Affect the Rate of Bioenergetics 124 Hydrolysis and Condensation: The Basis for Digestion and Synthesis 128

CHAPTER 6

Energy Transfer in the Body 134

Vitamins, Minerals, and Water 42 Part 1 • VITAMINS 43 The Nature of Vitamins 43 Kinds of Vitamins 43 Role of Vitamins 43 Defining Nutrient Needs 45 Exercise, Free Radicals, and Antioxidants 51 Vitamin Supplements: The Competitive Exercise Edge? 53

Optimal Nutrition for Exercise 81 Nutrient Intake Among the Physically Active 82 MyPyramid: The Essentials of Good Nutrition 87 Exercise and Food Intake 90 Precompetition Meal 94 Carbohydrate Feedings Prior to, During, and in Recovery from Exercise 97 Glucose Feedings, Electrolytes, and Water Uptake 101

Part 1 • PHOSPHATE BOND ENERGY 135 Adenosine Triphosphate: The Energy Currency 135 Phosphocreatine: The Energy Reservoir 138 Cellular Oxidation 138 Oxygen’s Role in Energy Metabolism 142 Part 2

• ENERGY RELEASE FROM

MACRONUTRIENTS 142 Energy Release from Carbohydrate 145 Energy Release from Fat 153 Energy Release from Protein 157 The Metabolic Mill: Interrelationships Among Carbohydrate, Fat, and Protein Metabolism 159

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CHAPTER 7

Energy Transfer During Exercise 162 Immediate Energy: The ATP–PCR System 163 Short-Term Energy: The Lactic Acid System 163 Long-Term Energy: The Aerobic System 164 Energy Spectrum of Exercise 168 Oxygen Consumption During Recovery 169

CHAPTER 8

Measurement of Human Energy Expenditure 178 Measuring the Body’s Heat Production 179 Doubly Labeled Water Technique 185 Respiratory Quotient 186 Respiratory Exchange Ratio 190

CHAPTER 9

Anatomy of Ventilation 254 Mechanics of Ventilation 255 Lung Volumes and Capacities 258 Lung Function, Aerobic Fitness, and Exercise Performance 261 Pulmonary Ventilation 263 Variations from Normal Breathing Patterns 265 The Respiratory Tract During Cold-Weather Exercise 266 CHAPTER 13

Part 1

Part 1 • ENERGY EXPENDITURE AT REST 193 Basal and Resting Metabolic Rate 193 Metabolic Size Concept 193 Comparing Metabolic Rates in Humans 194 Factors that Affect Energy Expenditure 197

CHAPTER 10

CHAPTER 11

Energy Expenditure During Walking, Jogging, Running, and Swimming 206 Gross Versus Net Energy Expenditure 207 Economy of Human Movement and Mechanical Efficiency 207 Energy Expenditure During Walking 209 Energy Expenditure During Running 212 Swimming 220 Individual Differences and Measurement of Energy Capacities 225 Specificity Versus Generality of Metabolic Capacity and Exercise Performance 226 Overview of Energy-Transfer Capacity During Exercise 226 Anaerobic Energy Transfer: The Immediate and Short-Term Energy Systems 227 Aerobic Energy: The Long-Term Energy System 234

Section 3

Part 3 • CARBON DIOXIDE TRANSPORT 282 Carbon Dioxide Transport in the Blood 282 CHAPTER 14

Pulmonary Structure and Function 253 Surface Area and Gas Exchange 254

Dynamics of Pulmonary Ventilation 286 Part 1

• REGULATION OF PULMONARY

VENTILATION 287 Ventilatory Control 287 Regulation of Ventilation During Exercise 289 Part 2

• PULMONARY VENTILATION DURING

EXERCISE 291 Ventilation and Energy Demands in Exercise 291 Energy Cost of Breathing 296 Does Ventilation Limit Aerobic Power and Endurance? 298 Part 3 • ACID–BASE REGULATION 300 Buffering 300 Physiologic Buffers 301 Effects of Intense Exercise 302 CHAPTER 15

The Cardiovascular System 303 Cardiovascular System Components 304 Hypertension 315 Blood Pressure Response to Exercise 317 The Heart’s Blood Supply 319 Myocardial Metabolism 322

CHAPTER 16

Cardiovascular Regulation and Integration 324 Intrinsic Regulation of Heart Rate 325 Extrinsic Regulation of Heart Rate and Circulation 328 Distribution of Blood 333 Integrative Exercise Response 335 Exercising after Cardiac Transplantation 335

CHAPTER 17

Functional Capacity of the Cardiovascular System 340 Cardiac Output 341 Cardiac Output at Rest 342

Aerobic Systems of Energy Delivery and Utilization 249

Interview with Dr. Loring B. Rowell CHAPTER 12

Part 2 • OXYGEN TRANSPORT 275 Transport of Oxygen in the Blood 275

• ENERGY EXPENDITURE DURING

PHYSICAL ACTIVITY 200 Classification of Physical Activities by Energy Expenditure 200 The MET 200 Daily Rates of Average Energy Expenditure 201 Energy Cost of Household, Industrial, and Recreational Activities 201 Heart Rate to Estimate Energy Expenditure 203

• GASEOUS EXCHANGE IN THE LUNGS

AND TISSUES 271 Concentrations and Partial Pressures of Respired Gases 271 Movement of Gas in Air and Fluids 272 Gas Exchange in the Lungs and Tissues 273

Human Energy Expenditure During Rest and Physical Activity 192

Part 2

Gas Exchange and Transport 270

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Cardiac Output During Exercise 343 Cardiac Output Distribution 346 Cardiac Output and Oxygen Transport 347 Cardiovascular Adjustments to Upper-Body Exercise 351 CHAPTER 18

Skeletal Muscle: Structure and Function 353 Gross Structure of Skeletal Muscle 354 Skeletal Muscle Ultrastructure 357 Muscle Fiber Alignment 360 Actin–Myosin Orientation 363 Chemical and Mechanical Events During Muscle Action and Relaxation 364 Muscle Fiber Type 371 Genes that Define Skeletal Muscle Phenotype 374 Fiber Type Differences Among Athletic Groups 374

CHAPTER 19

Neural Control of Human Movement 376 Neuromotor System Organization 377 Nerve Supply to Muscle 385 Motor Unit Functional Characteristics 390 Receptors in Muscles, Joints, and Tendons: The Proprioceptors 393

CHAPTER 20

The Endocrine System: Organization and Acute and Chronic Responses to Exercise 400 Endocrine System Overview 401 Endocrine System Organization 401 Resting and Exercise-Induced Endocrine Secretions 407 Gonadal Hormones 417 Exercise Training and Endocrine Function 430 Resistance Training and Endocrine Function 437 Opioid Peptides and Physical Activity 439 Physical Activity, Infectious Illness, Cancer, and Immune Response 439

American College of Sports Medicine and American Heart Association Updated Fitness Guidelines and Recommendations 475 How Long Before Improvements Occur? 476 Maintenance of Aerobic Fitness Gains 478 Training Methods 479 Overtraining: Too Much of a Good Thing 483 Exercising During Pregnancy 485 CHAPTER 22

Muscular Strength: Training Muscles to Become Stronger 490 Part 1

Section 4

Enhancement of Energy Transfer Capacity 447

Part 2

Training for Anaerobic and Aerobic Power 451 Exercise Training Principles 452 Physiologic Consequences of Exercise Training 457 Anaerobic System Changes with Training 458 Aerobic System Changes with Training 458 Factors that Affect Aerobic Training Responses 470

• STRUCTURAL AND FUNCTIONAL

ADAPTATIONS TO RESISTANCE TRAINING 519 Factors that Modify the Expression of Human Strength 519 Comparative Training Responses in Men and Women 526 Detraining 527 Metabolic Stress of Resistance Training 527 Circuit Resistance Training 528 Muscle Soreness and Stiffness 528 CHAPTER 23

Special Aids to Exercise Training and Performances 533 An Increasing Challenge to Fair Competition 534 A Need to Critically Evaluate the Scientific Evidence 538 On the Horizon 540 Pharmacologic Agents 541 Nonpharmacologic Approaches 566

Exercise Performance and Environmental Stress 587

Interview with Dr. Barbara Drinkwater CHAPTER 24

Exercise at Medium and High Altitude 591 The Stress of Altitude 592 Acclimatization 596 Metabolic, Physiologic, and Exercise Capacities at Altitude 604 Altitude Training and Sea-Level Performance 606 Combine Altitude Stay with Low-Altitude Training 608

CHAPTER 25

Exercise and Thermal Stress 611

Interview with Dr. Bengt Saltin CHAPTER 21

• STRENGTH MEASUREMENT AND

RESISTANCE TRAINING 491 Measurement of Muscle Strength 492 Gender Differences in Muscle Strength 495 Training Muscles to Become Stronger 498

Section 5

PART TWO: APPLIED EXERCISE PHYSIOLOGY 445

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Part 1

• MECHANISMS OF

THERMOREGULATION 612 Thermal Balance 612 Hypothalamic Temperature Regulation 613

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Contents Thermoregulation in Cold Stress: Heat Conservation and Heat Production 613 Thermoregulation in Heat Stress: Heat Loss 614 Effects of Clothing on Thermoregulation 618 Part 2

• THERMOREGULATION AND

ENVIRONMENTAL HEAT STRESS DURING EXERCISE 624 Exercise in the Heat 624 Maintaining Fluid Balance: Rehydration and Hyperhydration 627 Factors that Modify Heat Tolerance 630 Complications from Excessive Heat Stress 633 Part 3

Composition of the Human Body 733 Common Techniques to Assess Body Composition 738 Average Percentage Body Fat 756 Determining Goal Body Weight 757 CHAPTER 29

Physique, Performance, and Physical Activity 759 Physiques of Champion Athletes 760 Upper Limit for Fat-Free Body Mass 778

CHAPTER 30

Overweight, Obesity, and Weight Control 780 Part 1 • OBESITY 781 Historical Perspective 781 Obesity Remains a Worldwide Epidemic 781 A Progressive Long-Term Process 784 Genetics Influences Body Fat Accumulation 787 Physical Inactivity: A Crucial Component in Excessive Fat Accumulation 789 Health Risks of Excessive Body Fat 790 Criteria for Excessive Body Fat: How Fat Is Too Fat? 793

• THERMOREGULATION AND

ENVIRONMENTAL COLD STRESS DURING EXERCISE 635 Exercise in the Cold 635 Acclimatization to Cold 637 How Cold Is Too Cold? 638 CHAPTER 26

CHAPTER 27

Sport Diving 640 Diving History—Antiquity to the Present 641 Pressure–Volume Relationships and Diving Depth 645 Snorkeling and Breath-Hold Diving 646 Scuba Diving 650 Special Problems with Breathing Gases at High Pressures 653 Dives to Exceptional Depths: Mixed-Gas Diving 659 Energy Cost of Underwater Swimming 663 Microgravity: The Last Frontier 665 The Weightless Environment 666 Historical Overview of Aerospace Physiology and Medicine 671 Modern Era 673 Medical Evaluation for Astronaut Selection 673 Physiologic Adaptations to Microgravity 676 Countermeasure Strategies 693 Overview of Physiologic Responses to Spaceflight 709 Vision for the Future of Space Exploration 710 Practical Benefits from Space Biology Research 714

Section 6

Body Composition, Energy Balance, and Weight Control 721

Interview with Dr. Claude Bouchard CHAPTER 28

Body Composition Assessment 725 Overweight, Overfatness, and Obesity: No Unanimity for Terminology 728 The Body Mass Index: A Popular Clinical Standard 728

Part 2

• PRINCIPLES OF WEIGHT CONTROL: DIET

AND EXERCISE 801 Energy Balance: Input Versus Output 801 Dieting for Weight Control 802 Factors that Affect Weight Loss 810 Exercise for Weight Control 812 Effectiveness of Regular Physical Activity 814 Weight Loss Recommendations for Wrestlers and Other Power Athletes 822 Gaining Weight: The Competitive Athlete’s Dilemma 823

Section 7

Exercise, Successful Aging, and Disease Prevention 827

Interview with Dr. Steven N. Blair CHAPTER 31

Physical Activity, Health, and Aging 831 The Graying of America 832 Part 1

• PHYSICAL ACTIVITY IN THE

POPULATION 835 Physical Activity Epidemiology 835 Part 2

• AGING AND PHYSIOLOGIC FUNCTION

842 Age Trends 842 Trainability and Age 852 Part 3

• PHYSICAL ACTIVITY, HEALTH, AND

LONGEVITY 853 Causes of Death in the United States 854 Exercise, Health, and Longevity 854 Regular Moderate Exercise Provides Significant Benefits 856

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Contents

Can Increasing Physical Activity Level Improve Health and Extend Life? 859

Neuromuscular Diseases, Disabilities, and Disorders 919 Renal Disease 921 Cognitive/Emotional Diseases and Disorders 922

Part 4 • CORONARY HEART DISEASE 860 Changes on the Cellular Level 860 Coronary Heart Disease Risk Factors 864 CHAPTER 32

Clinical Exercise Physiology for Cancer, Cardiovascular, and Pulmonary Rehabilitation 876 The Exercise Physiologist in the Clinical Setting 877 Training and Certification Programs for Professional Exercise Physiologists 877 Clinical Applications of Exercise Physiology to Diverse Diseases and Disorders 879 Oncology 879 Cardiovascular Disease 885 Assessing Cardiac Disease 892 Stress Test Protocols 902 Prescribing Physical Activity and Exercise 904 Cardiac Rehabilitation 906 Pulmonary Diseases 909 Exercise and Asthma 917

xv

On the Horizon 929 Interview with Dr. Frank W. Booth

Molecular Biology—A New Vista for Exercise Physiology 933 Brief History Tour of Molecular Biology 936 Revolution in the Biologic Sciences 938 Human Genome 940 Nucleic Acids 942 How DNA Replicates 951 Protein Synthesis: Transcription and Translation 954 Mutations 971 New Horizons in Molecular Biology 979 Human Performance Research 1001 INDEX

1008

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USER’S GUIDE

Exercise Physiology, Seventh Edition, offers comprehensive coverage of exercise physiology uniting the topics of physical conditioning, sports nutrition, body composition, weight control, and more. To help your comprehension of the material, the authors have included numerous features that reinforce concepts and enhance your learning experience. Look for these features…

CHAPTER OBJECTIVES help you identify learning goals.

IN A PRACTICAL SENSE connects theoretical concepts to practical skills.

FOCUS ON RESEARCH presents a key research article from a renowned scientist and illustrates how “theory comes to life.”

INTEGRATIVE QUESTIONS pose open-ended questions for reflection on complex concepts.

Animations, quizzes, appendices, and other student resources found on thePoint at http://thepoint.lww.com/mkk7e

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INTRODUCTION: A V I E W O F T H E PA S T

EXERCISE PHYSIOLOGY: ROOTS AND HISTORICAL PERSPECTIVES Since the first edition of our textbook 28 years ago in 1981, knowledge has exploded concerning the physiologic effects of exercise in general, and the body’s unique and specific responses to training in particular. Tipton’s search of the 1946 English literature for the terms exercise and exertion yielded 12 citations in 5 journals.66 Tipton also cited a 1984 analysis by Booth, who reported that in 1962, the number of yearly citations of the term exertion increased to 128 in 51 journals, and by 1981, there were 655 citations for the word exertion in 224 journals. The accompanying figure reveals the number of entries for the words exercise or exertion from an Internet search of Index Medicus (Medline) for the years 2000 to December 18, 2008, using the National Center for Biotechnology Information (NCBI) database (www.ncbi.nlm.nih.gov/ sites/entrez). In just a 4-year period since publication of the sixth edition, the number of citations increased tremendously to 224,421 (312%)! For the fourth edition published in 1996,

230,000 224,421

225,000 220,000 60,000

57,180

55,000 50,829

Number of citations

50,000 45,000

43,625

40,000 4000 3500 3000 2500

2154

2069

1288

1441

1986

1996 2000 Year

2000 1500 1249

1000 500 0

703 342

1966

540

1976

2003

2005

2009

Exercise or exertion as a topic (top bars) and frequency of the word exercise appearing in a scientific journal (bottom bars) for the years 1966 to 1996 (from Index Medicus). The last three columns used PubMed via an Internet search for citations with the terms exercise or exertion.

we noted that the greatest increases occurred between 1976 and 1986, and that citation frequency appeared to level off from 1986 and 1996. From that time, the rate of increase has been even greater. Obviously, we misjudged how greatly exercise-related topics would affect scholarly productivity in biologic sciences research. The number of citations has increased beyond our wildest expectations, and with expanding interest in the role of exercise and physical activity, the rate of citations devoted to these topics undoubtedly will continue to accelerate. As graduate students in the late 1960s, we never believed that interest in exercise physiology would increase so dramatically. A new generation of scholars committed to studying the scientific basis of exercise had set to work. Some studied the physiologic mechanisms involved in adaptations to regular exercise; others evaluated individual differences in exercise and sports performance. Collectively, both approaches contributed knowledge to the growing field of exercise physiology. At our first scientific conference (American College of Sports Medicine [ACSM] in Las Vegas, 1967), still as graduate students, we rubbed elbows with the “giants” of the field, many of whom were themselves students of the leaders of their era. Sitting under an open tent in the Nevada desert with one of the world’s leading physiologists, Dr. David Bruce Dill (then age 74; see page xlvii), we listened to his researcher—a high school student—lecture about temperature regulation in the desert burro. Later, one of us (FK) sat next to a whitehaired gentleman and chatted about a Master’s thesis project. Only later did an embarrassed FK learn that this gentleman was Captain Albert R. Behnke, MD (1898–1993; ACSM Honor Award, 1976), the modern-day “father” of human body composition assessment whose crucial experiment in diving physiology established standards for decompression and use of mixed gases. His pioneering studies of hydrostatic weighing in 1942, the development of a reference man and reference woman model, and creation of the somatogram based on anthropometric measurements underlie much current work in body composition assessment (see Chapter 28 and its “Focus on Research”). That fortuitous meeting began a lasting personal and fulfilling professional friendship until Dr. Behnke’s death in 1993. Several hundred ACSM members listened attentively as the superstars of exercise physiology and physical Albert R. Behnke xvii

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Introduction: A View of the Past

fitness (Per-Olof Åstrand, Erling Asmussen, Bruno Balke, Elsworth Buskirk, Thomas Cureton, Lars Hermansen, Steven Horvath, Henry Montoye, Bengt Saltin, Charles Tipton) presented their research and fielded penetrating questions from an audience of young graduate students eager to devour the latest scientific information. Over the years, the three of us have been fortunate to work with the very best in our field. William McArdle studied for the PhD at the University of Michigan with Dr. Henry Montoye (charter member of ACSM; President of ACSM, 1962–1963; Citation Award, 1973) and Dr. John Faulkner (President of ACSM, 1971–1972; Citation Award, 1973; and ACSM Honor Award, 1992). At the University of California, Berkeley, Victor Katch completed the MS thesis in physical education under the supervision of Dr. Jack Wilmore (ACSM President, 1978–1979; Citation Award, 1984; and first editor of Exercise and Sport Science Reviews, 1973–1974) and was a doctoral student of Dr. Franklin Henry (ACSM Honor Award, 1975; originator of the “Memory-Drum Concept” about the specificity of exercise; and author of the seminal paper “Physical Education—an Academic Discipline,” JOHPER 1964;35:32). Frank Katch completed the MS degree at the University of California, Santa Barbara, under the supervision of thesis advisors Dr. Ernest Michael, Jr., (former PhD student of pioneer exercise physiologist–physical fitness scientist Dr. Thomas Kirk Cureton, ACSM Honor Award, 1969) and Dr. Barbara Drinkwater (President of ACSM, 1988–1989; ACSM Honor Award, 1996), and then also completed doctoral studies at the University of California, Berkeley, with Professor Henry. Professor Roberta Park, also at the University of California, Berkeley, was inspirational over the years in sparking interest in the history of physical education as an academic discipline. She encouraged each of us (and our future students) to honor the past accomplishments of those in the profession who helped to pave the way. As the three of us examine those earlier times, we realize, like many of our colleagues, that our academic good fortunes prospered because our professors and mentors shared an unwavering commitment to study sport and exercise from a strong scientific and physiologic perspective. These scholars demonstrated why it was crucial that physical educators be well grounded in both the scientific basics and underlying concepts and principles of exercise physiology. We are so pleased to acknowledge the pioneers who created exercise physiology, realizing full well the difficult task in an introduction to adequately chronicle the history of exercise physiology from its origins in ancient Asia to the present. Instead, our brief review presents a historical tour regarding topics not adequately developed in exercise physiology or history textbooks. Our discussion begins with an acknowledgment of the ancient but tremendously influential Indian, Arabic, and prominent Greek physicians; along the way, we highlight some milestones (and ingenious experiments), including the many contributions from Sweden, Denmark, a

Norway, and Finland that fostered the study of sport and exercise as a respectable field of scientific inquiry. A treasure of information about the early beginnings of exercise physiology in America was uncovered in the archives of Amherst College, Massachusetts, in an anatomy and physiology textbook (incorporating a student study guide) written by the first American father-and-son writing team. The father, Edward Hitchcock, was President of Amherst College; the son, Edward Hitchcock, Jr., an Amherst graduate and Harvard-trained physician, made detailed anthropometric and strength measurements of almost every student enrolled at Amherst College from 1861 to 1889. A few years later in 1891, much of what forms current college curricula in exercise physiology, including evaluation of body composition by anthropometry and muscular strength by dynamic measurements, began in the first physical education scientific laboratory at Harvard University’s Lawrence Scientific School. Even before the creation of this laboratory, another less formal but still tremendously influential factor affected the development of exercise physiology: the publication during the 19th century of American textbooks on anatomy and physiology, physiology, physiology and hygiene, and anthropometry. TABLE 1 lists a sampling of textbooks published between 1801 and 1899 containing information about the muscular, circulatory, respiratory, nervous, and digestive systems—including the influence of exercise and its effects— that eventually shaped the content area of exercise physiology during the next century. Roberta Park, distinguished UC Berkeley physical education historian, chronicles the early contributions of many physicians and science-oriented physical educators who steadfastly believed that physical education (and medicine) should be grounded on a sound scientific foundation fueled by cutting-edge research.50,52,54,56 These well-documented historical chronologies and other contributions51,53,55 provide context and foster appreciation for the scholars and educators who paved the way for the new generation of researchers; the early innovators developed new techniques and methodologies in the fields of health, fitness, performance, and physical activity that became essential components of the exercise physiology core curriculum. Textbooks from 1900 to 1947 dealing with exercise, training, and exercise physiology also were influential.a

IN THE BEGINNING: ORIGINS OF EXERCISE PHYSIOLOGY FROM ANCIENT GREECE TO AMERICA IN THE EARLY 1800s Exercise physiology arose mainly in early Greece and Asia Minor, although the topics of exercise, sports, games, and health concerned even earlier civilizations. These included the Minoan and Mycenaean cultures, the great biblical empires of David and Solomon, Assyria, Babylonia, Media,

Buskirk11 provides a bibliography of books and review articles on exercise, fitness, and exer