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 EIGHTH EDITION

Exercise
Physiology
Nutrition, Energy, and Human Performance

McArdle9781451191554_FM.indd 1 12/11/2013 7:50:44 PM

McArdle9781451191554_FM.indd 2 12/11/2013 7:50:44 PM
 EIGHTH EDITION

Exercise
Physiology
Nutrition, Energy, and Human Performance

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

Frank I. Katch
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 of Movement Science
School of Kinesiology
Associate Professor, Pediatrics
School of Medicine
University of Michigan, Ann Arbor, Michigan

McArdle9781451191554_FM.indd 3 12/11/2013 7:50:49 PM

Acquisitions Editor: Emily Lupash
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Marketing Manager: Shauna Kelley
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Copyright © 2015, 2010, 2007, 2001, 1996, 1986, 1981

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Selected photographs © 2008 by Fitness Technologies, Inc., Frank I. Katch, and Victor L. Katch. This mate-
rial is protected by copyright. No photographs may be reproduced in any form or by any means without
permission from the copyright holders.

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9 8 7 6 5 4 3 2 1
 To my wife Kathleen, my best friend and biggest supporter, and to the rest of the
“A team,” whose lives give meaning to my own: my children, Theresa, Amy, Kevin,
and Jennifer; their spouses, Christian, Jeff, Nicole, and Andy;
and my grandchildren, Liam, Aidan, Dylan, Kelly Rose, Owen,
Henry, Kathleen (Kate), Grace, Elizabeth, Claire, Elise,
Charlotte, and Sophia.

—Bill McArdle

To my wife and life partner Kerry for 44 years of love, patience, and support;
to my two sons, David and Kevin, for achieving the honorable in their
professional lives; to my daughter Ellen (and her husband Sean) for all
her success as a caring pediatrician and mom; and to my
one-year-old grandson pal, James Patrick. Life is good!

— F r a n k K at c h

To those most important to me: my wife Heather, my daughters

Erika and Leslie, my son Jesse, and my grandkids
Ryan, Cameron, Ella, and Emery.

— V i c t o r K at c h

McArdle9781451191554_FM.indd 5 12/11/2013 7:50:49 PM

 Preface

Since the first edition of our textbook more than threee decades health professions, the rate of citations devoted to these topics
ago, knowledge concerning the physiologic effects of exercise in undoubtedly will continue to accelerate.
general and the body’s unique and specific responses to training As graduate students in the late 1960s, we never imagined
in particular has exploded. Tipton’s search of the 1946 English that interest in exercise physiology would increase so dramati-
literature for the terms exercise and exertion yielded 12 citations cally. New generations of scholars committed to studying the
in 5 journals.73 Tipton also cited a 1984 analysis by Booth, who scientific basis of exercise had set to work. Some studied the
reported that in 1962, the number of yearly citations of the term physiologic mechanisms involved in adaptations to regular
exertion increased to 128 in 51 journals, and by 1981, there were exercise; others evaluated individual differences in exercise and
655 citations to the word exertion in 224 journals. The graph on sports performance. Collectively, both approaches expanded
this page highlights the huge number of entries for the words knowledge in the growing field of exercise physiology. At our
exercise or exertion from a recent Internet search of Index Medicus first scientific conference (American College of Sports Medi-
(Medline) and for the years 2000 to December 3, 2013, using the cine [ACSM] in Las Vegas, 1967), still as graduate students, we
NCBI database (www.ncbi.nlm.nih.gov/sites/entrez). In just a rubbed elbows with the “giants” of the field, many of whom were
4-year period since publication of our seventh edition, the num- themselves students of the leaders of their era. Several hundred
ber of citations increased by over 66,700 to 291,194, a 29.8% ACSM members listened attentively as the superstars of exer-
increase! Although we had thought that citation frequency was cise physiology and physical fitness (Erling Asmussen, Per-Olof
leveling off from 1986 to 1996, the rate of increase has instead Åstrand, Bruno Balke, Elsworth Buskirk, Thomas Cureton,
steadily increased beyond our wildest expectations. Obviously, we Lars Hermansen, Steven Horvath, Henry Montoye, Bengt
misjudged how greatly exercise-related topics would affect schol- Saltin, and Charles Tipton) presented their research and fielded
arly productivity in biologic sciences research. With expanding penetrating questions from an audience of young graduate stu-
interest in the role of exercise and physical activity in the allied dents eager to devour the latest scientific information delivered
by these “stars of our field.”
Sitting under an open tent in the Nevada
desert with one of the world’s leading physiolo-
290,000
288,379 gists, Dr. David Bruce Dill (then age 74; profiled
later in this book’s introduction), we listened to
285,000
his research assistant—a high school student—
280,000 lecture about temperature regulation in the des-
230,000 ert burro. Later, one of us (Frank Katch) sat next
225,000 224,421 to a white-haired gentleman and chatted about
his master’s thesis project. Only later did an
220,000
embarrassed Frank learn that this gentleman was
60,000
57,180
Captain Albert R. Behnke, MD (1898–1993;
ACSM Honor Award, 1976), the modern-day
Number of citations

55,000
50,829 “father” of human body composition assessment
50,000
whose crucial experiment in diving physiology
45,000
43,625
established standards for decompression and use
40,000 of mixed gases for deep dives.
Dr. Behnke’s pioneering studies of hydro-
4000
static weighing in 1942 (which Frank Katch put
3500
into practice with a swimming pool underwater
3000 weighing tank for his master of science thesis at the
2500
University of California, Santa Barbara, in 1966),
2069
2154 the development of a reference man and reference
2000
woman model, and the creation of the somato-
1500 gram based on anthropometric measurements
1249
1000
703
Exercise or exertion as a topic (top bars) and
500 1288 1441 frequency of the word exercise appearing in
540
0
342 a scientific journal (bottom bars) for the years
1966 1976 1986 1996 2000 2003 2005 2009 2013 1966 to 2013 from Index Medicus. The last four
Year columns used PubMed via an Internet search
for citations with the terms exercise or exertion.
vi

McArdle9781451191554_FM.indd 6 12/11/2013 7:50:51 PM

 Preface vii

form the basis for much cur- exercise. As an example, proper nutrition links to good health,
rent work in body composi- effective weight control, and optimal levels of physical activ-
tion assessment. ity and sports performance, while regular physical activity
That fortuitous meeting and exercise training provide an important means to control
began a lasting personal and body weight and optimize one’s overall health profile. We are
fulfilling professional friend- encouraged that the medical establishment and government
ship until Dr. Behnke’s death agencies continue to acknowledge (and now promote) regular
in 1993. Over the years, the physical activity as an important weapon in the armamentar-
three of us were indeed for- ium for prevention and rehabilitation of diverse disease states,
tunate to work with the very including diabetes, obesity, cancer, and heart disease.
best scholars in our field. We are gratified with the small part we have played in the
William McArdle studied education of more than 400,000 undergraduate and graduate
for his PhD at the University students who have used this text since the publication of the
of Michigan with Dr. Henry first edition in 1981. A source of great pride for us is that some
Albert R. Behnke
Montoye (charter member of the first students enrolled in our classes that used this text
of ACSM; President of ACSM, 1962–1963; Citation Award, have gone on to earn advanced degrees in the same or simi-
1973) and Dr. John Faulkner (President of ACSM, 1971–1972; lar fields. This tradition of textbook adoption has now been
Citation Award, 1973; ACSM Honor Award, 1992). At the passed down to their students, many of whom comprise the
University of California, Berkeley, Victor Katch completed next generation of aspiring teachers, exercise specialists, and
his master of science thesis under the supervision of Dr. Jack researchers. We are forever grateful to our former teachers and
Wilmore (ACSM President, 1978–1979; Citation Award, 1984; mentors for igniting a spark that has not diminished. We hope
first editor of Exercise and Sport Science Reviews, 1973–1974) and you will become as excited as we first were (and continue to be)
was a doctoral student of Dr. Franklin Henry (ACSM Honor in the science of exercise physiology and human performance.
Award, 1975; originator of the “Memory-Drum Concept” about We leave you with this apt quote in Latin attributed to pro-
the specificity of exercise; author of the seminal paper “Physical lific French author and astronomer Nicolas Camille Flammarion
Education—an Academic Discipline,” JOHPER 1964;35:32). (1842 –1925): “Ad Veritatum Per Scientiam” (To Truth Through
Frank Katch completed his master of science under the super- Science), inscribed in gold above the observatory and museum
vision of thesis advisors Dr. Ernest Michael, Jr. (former PhD entrance to his Chateau at Juvisy-Sur-Orge outside of Paris.
student of pioneer exercise physiologist–physical fitness scien-
tist Dr. Thomas Kirk Cureton; ACSM Honor Award, 1969), ORGANIZATION
and Dr. Barbara Drinkwater (President of ACSM, 1988–1989;
ACSM Honor Award, 1996), and then completed doctoral This eighth edition maintains an eight-section structure and an
studies at UC Berkeley with Professor Franklin Henry. introductory section about the origins of exercise physiology.
As the three of us reexamine those earlier times, we real- The concluding “On the Horizons” section and its chapter have
ize, like many of our colleagues, that our academic good for- changed from an addendum to a numbered chapter, reflective of
tunes prospered because our professors and mentors shared an molecular biology’s place as an established part of exercise science.
unwavering commitment to study sport, exercise, and move- The eighth edition also has undergone a complete art
ment from a strong scientific and physiologic perspective. makeover. Most of the existing figures have been redrawn to
These scholars demonstrated why it was crucial that physical provide consistency with newly created illustrations. Through-
educators be well grounded in both the scientific basics and out the text, we have included Internet resources (URLs)
underlying concepts and principles of exercise physiology. to provide an expanded Web access to supplement student
insights of relevant text material. The text continues the tradi-
tion of FYI (For Your Information) boxes that provide rela-
Moving Forward tively short inserts of related information, current research, or
As in the publication of the first edition of Exercise Physiol- interesting sidebars germane to the text’s topic, ranging from
ogy: Energy, Nutrition, and Human Performance in 1981, this “One-Minute Bouts of Intense Physical Activity Improves
eighth edition reflects our continued commitment to inte- Fitness and Health” to “Consuming Excess Calories Produces
grate the concepts and science of the different disciplines Fat Gain Regardless of Nutrient Source.”
that contribute to a more comprehensive understanding and
appreciation of modern-day exercise physiology. As in prior FEATURES
editions, we continue to believe that the exercise physiol-
This text’s features have been specifically designed to help stu-
ogy discipline demands integration of study areas relevant to
dents facilitate learning. They include:
nutrition, exercise biochemistry and bioenergetics, physiology,
medicine, exercise training and sports performance, and the Introduction: A View of the Past. The text’s introduction,
health-related aspects of regular physical activity. All of these “Exercise Physiology: Roots and Historical Perspectives,”
areas naturally and inexorably link within the fabric of what ­reflects our interest and respect for the earliest underpinnings
currently comprises the field called exercise physiology or, in of the field, and the direct and indirect contributions of the men
deference to the early scholars in the field, the physiology of and women physicians–scientists who contributed to the field.

McArdle9781451191554_FM.indd 7 12/11/2013 7:50:51 PM

 viii Preface

Chapter Objectives. Each chapter opens with a comprehen- to reflect current research findings related to the diverse areas
sive summary of learning goals, helping students to become of exercise physiology. We have revised almost every figure,
familiar with the materials to be covered in a chapter. and supplemented them with high-quality medical illustra-
Ancillaries at-a-Glance. A complete list of all electronic tions. We also have added new tables, and listed numerous
­resources associated with a chapter makes accessing online new Web sites to provide readers access to the abundance of
­materials easy; callouts in the text reinforce for students oppor- updated information available about the intricacies relevant to
tunities to broaden their knowledge beyond the pages of the text. topic areas in exercise physiology. “On the Horizon” has been
upgraded to a full section and chapter, reflecting the increas-
In a Practical Sense. Every chapter highlights practical
ing importance of research in molecular biology on exercise
­applications about specific topic areas.
physiology.
Integrative Questions. Open-ended questions encourage Our current reference list includes up-to-date research
students to thoughtfully consider complex concepts without results gleaned from national and international journals
a single “correct” answer. related to specific topic areas. In selected chapters, “Additional
Expanded Art Program. The full-color art program continues References” provide a bibliography of articles that augment
to be an important feature of the textbook. Nearly every figure the materials already presented in the chapter. All references
has been revised to make its textual and visual elements “pop,” or for a chapter are located online at http://thePoint.lww.com/
altered to highlight important teaching points that reinforce text mkk8e. We hope you profit from and enjoy this continuation
material. New figures have been added to chapters to enhance of our journey through the ever-expanding and maturing field
new and updated content, including the use of many new medical of exercise physiology.
illustrations. A new table format clearly organizes essential data.
Up-Close and Personal Interviews. The text features nine ANCILLARIES: THE TOTAL TEACHING
contemporary scientists whose important research contributions PACKAGE
and visionary leadership continue the tradition of the scientists
of prior generations—Drs. Steven Blair, Frank Booth, Claude Exercise Physiology: Nutrition, Energy, and Human Performance,
Bouchard, David Costill, Barbara Drinkwater, John Holloszy, Eighth Edition includes additional resources for both instruc-
Loring Rowell, Bengt Saltin, and Charles Tipton. These individ- tors and students that are available on the book’s companion
uals merit recognition not only for expanding knowledge through Web sites at http://thePoint.lww.com/mkk8e.
their many scientific contributions, but also for elucidating mech- Approved adopting instructors will be given access to the
anisms that underlie responses and adaptations to exercise and following resources:
health enhancement. Each person has been placed within a sec- ●● Animations illustrating the most important concepts in
tion linked to his or her main scholarship interests, yet all of them human physiology
span one or more sections in terms of scientific contributions. ●● Test generator
Appendix C, available online at http://thepoint.
●● PowerPoint presentations: one set with lecture outlines,
lww.com/mkk8e, lists individual honors and one set with images only
awards for each of these distinguished and
●● Image bank of downloadable figures and tables in multiple
meritorious scientist–researchers. formats
●● Searchable full text online
The intimate insights from the “superstars” should inspire ●● Blackboard, Angel, and Moodle LMS cartridges
current exercise physiology students to actualize their poten-
tial, whether through accomplishments in graduate school, Students
teaching, research, or numerous other exciting professional Students who purchase Exercise Physiology: Nutrition, Energy,
opportunities to achieve excellence. and Human Performance, Eighth Edition have access to the
References, Appendices, and Animations (available following additional resources, accessible with the scratch-off
online). All references and appendices are available online at code provided on this book’s inside cover:
http://thepoint.lww.com/mkk8e. Appendices feature valu-
●● Online interactive quiz bank with study and test options
able information about nutritive values, energy expenditures,
●● Animations
metabolic computations in open-circuit spirometry, and more.
●● References
Focus on Research (available online). Almost all chap- ●● Appendixes
ters have a companion online Focus on Research, featuring ●● Focus on Research article abstracts and analysis
a key research article from a renowned scientist. These well- ●● Featured information on microscope technologies, notable
designed studies illustrate, within a historical perspective, how events in genetics, Nobel prizes, outstanding female scien-
“theory comes to life” via the dynamics of research. tists, and much more.
Ancillaries were prepared by the authors and by Jeff Woods
NEW TO THE EIGHTH EDITION (Professor of Kinesiology and Community Health, University
The flow of information in this edition remains similar to prior of Illinois at Urbana–Champaign) and Lamia Scherzinger
editions. Components of the entire text have been upgraded (Indiana University–Perdue University Indianapolis).

McArdle9781451191554_FM.indd 8 12/11/2013 7:50:52 PM

 Acknowledgments

We wish to thank many individuals. First, to Dr. Loring history MI. Dr. Barry Franklin, Beaumont Hospital, Detroit,
Rowell for his constructive comments on the chapters related MI, supplied original information about cardiac rehabilita-
to pulmonary and cardiovascular dynamics during rest and tion. The Trustees of Amherst College and Archival Library,
exercise, particularly the sections related to the possible role Amherst, MA, gave permission to reproduce the photographs
of the venous system as an active vasculature. We thank and materials of Dr. Hitchcock. Magnus Mueller from the
Dr. Victor Convertino, U.S. Army Institute of Surgical University of Geisen, Germany, kindly provided the photo
Research at Fort Sam Houston, TX, for insightful comments of Liebig’s Geisen lab. We are grateful to marine artist Ron
and suggestions on the microgravity chapter, and Dr. Charles Scobie, ASMA (www.­ronscobie-marineartist.com), for his
­
Tipton, Professor Emeritus, University of Arizona, Tuscon, kind permission to reproduce his rendering of the HMS
AZ, for valuable comments and for providing new informa- Beagle. We thank Nancy Mullis for graciously providing the
tion about the historical development of the physiology of photo of Dr. Kary Mullis.
exercise, including material about the first textbook devoted We are collectively indebted to the nine researchers/
to exercise and physiology in the 16th century, and physiology scholars who took time from their busy schedules to answer
of exercise textbook used in the late 1800s and early 1900s. our interview questions and provide personal photos. Each
Stephen Lee (Exercise Physiology Laboratory, ­Johnson of those individuals, in his or her own unique way, inspired
Space Center, Houston, TX; www.nasa.gov/centers/johnson/ the three of us in our careers by their work ethic, scientific
slsd/about/divisions/hacd/laboratories/exercise-­physiology. excellence, and generosity of time and advice with colleagues
html) kindly supplied original NASA photos and documents, and students. Over the years, we have had the good fortune
and Mission Specialist Astronaut Dr. Martin Fettman (Col- to come to know these individuals both socially and in the
orado State University, Ft. Collins, CO) provided an origi- academic arena. We are grateful for the opportunity to con-
nal slide of the rotating chair experiment he took during his duct the interviews because they provided insights about their
Skylab 2 Mission. Dr. Helen Lane (Chief Nutritionist and personal lives previously unknown to us. We hope you too
Manager, University Research and Affairs, NASA ­Johnson are as impressed as we are by all they have accomplished and
Space Center, Houston, TX), provided prepublication docu- returned to the profession.
ments and resource materials. Dr. Ron White, National We acknowledge our master’s and senior honors students
Space Biomedical Research Institute Houston, TX, allowed who worked in our labs for their projects, and contributed so
us to use charts he helped to create from Human Physiology much to our research and personal experiences: Pedro Alexan-
in Space Teacher’s Manual. Dr. Susan Bloomfield (Bone Biol- der, Christos Balabinis, Margaret Ballantyne, Brandee Black,
ogy Laboratory, Texas A&M University, College Station, TX) Michael Carpenter, Steven Christos, Roman Czula, Gwyn
kindly provided images of hind-limb suspension experiments Danielson, Toni Denahan, Marty Dicker, Sadie Drumm, Peter
from her lab. We sincerely appreciate the expertise of Drs. Frykman, Scott Glickman, Marion Gurry, Carrie Hauser,
Frank Booth, University of Missouri, Columbia, MO; Kris- Margie King, Peter LaChance, Jean Lett, Maria Likomitrou,
tin Steumple, Department of Health and Exercise Science at Robert Martin, Cathi Moorehead, Susan Novitsky, Joan Perry,
Gettysburg College, Gettysburg, PA; and Marvin Balouyt, Sharon Purdy, Michelle Segar, Debra Spiak, Lorraine ­Turcotte,
Washtenaw Community College, Ann Arbor, MI, for their Lori Waiter, Stephen Westing, and Howard Zelaznik.
expert opinions and suggestions for improving the chapter We also dedicate this edition to that special group of for-
on molecular biology. Hypoxico Inc. provided photos of the mer students who earner doctoral degrees in physical educa-
Hypoxico altitude tent. Mr. John Selby (www.hyperlite.co.uk) tion, exercise science, or medicine, and who have gone on to
kindly provided timely information and photos of the porta- distingush themselves as teachers, practitioners, and research-
ble, collapsible decompression chamber. Dr. Alex Knight, York ers in the related areas of exercise physiology. These include
University, United Kingdom, graciously provided information Denise Agin, Stamitis Agiovlasitis, Doug Ballor, Dan Becque,
about molecular biology techniques he has pioneered (in vitro Geroge Brooks, Barbara Campaigne, Ed Chaloupka, Ken
motility assay) and other information and a photograph about Cohen, Edward Coyle, Dan Delio, Julia Chase Delio, Chris
myosin, muscle, and single molecules. Yakl Freedman (www. Dunbar, Patti Freedson, Roger Glaser, Ellen Glickman, Kati
dna2z.com) was supportive in supplying recent information Haltiwinger, Everett Harmon, Jay Hoffman, Tibor Hortoba-
about DNA and molecular biology. Sue Hilt of the American gyi, Jie Kang, Mitch Kanter, Betsy Keller, Marliese Kimmerly,
College of Sports Medicine, Indianapolis, IN, did a superb George Lesmses, Steve Lichtman, Charles Marks, Robert
job of securing the text of the Citation and Honor Awards Mofatt, Laren Nau-White, Steve Ostrove, James Rimmer,
reproduced in Appendix C. Dr. James A. Freeman, Professor Deborah Rinaldi, Stan Sady, Lapros Sidossis, Bob Spina, John
of English, University of Massachusetts, Amherst, unselfishly Spring, Bill Thorland, Mike Toner, Laurel Trager-Mackin-
lent his know-how to make words sing in the introductory non, Lorraine Turcotte, John Villanacci, Jonnis Vrabis, Nancy

ix

McArdle9781451191554_FM.indd 9 12/11/2013 7:50:53 PM

 x Acknowledgments

Weiss, Art Weltman, Nancy Wessingeer, Stephen Westing, every chapter and dealing with our sometimes trivial requests.
Anthony Wilcox, and Libnda Zwiren. David Orzechowski, Production Project Manager, helped to
Finally, we would like to recognize the creative individuals translate the edited chapters into galley magic. Also, the tal-
at Wolters Kluwer who helped to shepherd this eighth edi- ented artists at Dragonfly (www.dragonflymediagroup.com/)
tion through the various stages of production. We are par- deserve recognition for their elegant medical illustration and
ticularly indebted to Eve Malakoff-Klein, our talented and artistic and technical expertise. Thank you so much Eve, Jen,
superb Supervisor of Product Development, who continu- Dave, and Dragonfly for a job exceptionally well done!
ally provided much-needed support, patience, subtle urging,
and excellence in organization and expertise in handling William D. McArdle
critical editing issues in bringing this edition to fruition in Sound Beach, NY
a timely manner. She clearly served in a highly professional Frank I. Katch
manner as our advocate in issues related to the production Santa Barbara, CA
process. We also gratefully acknowledge and appreciate the
Victor L. Katch
outstanding technical and creative expertise of Jennifer Cle-
Ann Arbor, MI
ments, Art Director, for going well beyond the call of duty
for insightful and creative contributions in revising the art in

McArdle9781451191554_FM.indd 10 12/11/2013 7:50:53 PM

 Contents

Introduction Water Balance: Intake Versus Output 74

A View of the Past: Exercise Physiology: Water Requirement in Physical Activity 75
Roots and Historical Perspectives xv Chapter 3
Interview with Dr. Charles M. Tipton Optimal Nutrition for Physical Activity 79
Nutrient Intake Among the Physically Active 80
The Essentials of Good Nutrition 85
Part One Myplate: The Healthy Eating Guide 85
Physical Activity and Food Intake 87
Exercise Physiology 1 Precompetition Meal 92
Carbohydrate Feedings Prior To, During, and in Recovery from
Section 1 Nutrition: The Base for Physical Activity 94
Glucose Feedings, Electrolytes, and Water Uptake 99
Human Performance 3
Interview with Dr. David L. Costill Section 2 Energy for Physical
Chapter 1 Activity105
Carbohydrates, Lipids, and Proteins 7 Interview with Dr. John O. Holloszy
PART 1 • CARBOHYDRATES 8 Chapter 4
Kinds and Sources of Carbohydrates 8 Energy Value of Food 109
Recommended Intake of Carbohydrates 14
Role of Carbohydrates in The Body 14 Measurement of Food Energy 110
Carbohydrate Dynamics During Physical Activity 15 Chapter 5
PART 2 • LIPIDS 18 Introduction to Energy Transfer 117
The Nature of Lipids 18
Kinds and Sources of Lipids 18 Energy—The Capacity for Work 118
Recommended Lipid Intake 25 Interconversions of Energy 120
Role of Lipid in The Body 26 Biologic Work in Humans 121
Fat Dynamics During Physical Activity 27 Enzymes and Coenzymes Alter the Rate of Energy Release 124
Hydrolysis and Condensation: The Basis for Digestion
PART 3 • PROTEINS 30
and Synthesis 126
The Nature of Proteins 30
Kinds of Protein 30 Chapter 6
Recommended Protein Intake 32 Energy Transfer in the Body 133
Protein’s Role in The Body 33
Dynamics of Protein Metabolism 33 PART 1 • PHOSPHATE BOND ENERGY 134
Nitrogen Balance 36 Adenosine Triphosphate: The Energy Currency 134
Protein Dynamics During Physical Activity 37 Phosphocreatine: The Energy Reservoir 136
Cellular Oxidation 137
Chapter 2 Oxygen’s Role in Energy Metabolism 140
Vitamins, Minerals, and Water 41 PART 2 • ENERGY RELEASE FROM MACRONUTRIENTS 141
PART 1 • VITAMINS 42 Energy Release from Carbohydrate 141
The Nature of Vitamins 42 Energy Release from Fat 150
Kinds of Vitamins 42 Energy Release from Protein 156
Role of Vitamins 42 The Metabolic Mill: Interrelationships Among Carbohydrate,
Defining Nutrient Needs 43 Fat, and Protein Metabolism 156
Physical Activity, Free Radicals, and Antioxidants 49 Chapter 7
Does Vitamin Supplementation Provide a Competitive Edge? 51 Energy Transfer During Physical Activity 161
PART 2 • MINERALS 53
The Nature of Minerals 53 Immediate Energy: The Atp–Pcr System 162
Role of Minerals in The Body 53 Short-Term Glycolytic (Lactate-Forming) Energy System 162
Calcium 57 Long-Term Energy: The Aerobic System 163
The Female Athlete Triad: An Unexpected Problem for Women Energy Spectrum of Physical Activity 167
Who Train Intensely 61 Oxygen Consumption During Recovery 168
Phosphorus 63 Chapter 8
Magnesium 63 Measurement of Human Energy Expenditure 177
Iron 64
Sodium, Potassium, and Chlorine 67 Measuring The Body’s Heat Production 178
Minerals and Exercise Performance 68 Doubly Labeled Water Technique 184
PART 3 • water 72 Respiratory Quotient 185
The Body’s Water Content 72 Respiratory Exchange Ratio (Rer) 189

xi

McArdle9781451191554_FM.indd 11 12/11/2013 7:50:53 PM

 xii Contents

Chapter 9 PART 3 • CARBON DIOXIDE TRANSPORT 282

Human Energy Expenditure During Rest Carbon Dioxide Transport in The Blood 282
and Physical Activity 191 Chapter 14
PART 1 • Energy Expenditure at Rest  192 Dynamics of Pulmonary Ventilation 285
Basal and Resting Metabolic Rate 192 PART 1 • REGULATION OF PULMONARY VENTILATION 286
Metabolic Size Concept 192
Ventilatory Control 286
Metabolic Rates of Humans: Age and Gender Comparisons 193
Regulation of Ventilation During Physical Activity 288
Five Factors That Affect Total Daily Energy Expenditure 196
PART 2 • P
 ULMONARY VENTILATION DURING PHYSICAL
PART 2 • Energy Expenditure during Physical
ACTIVITY 290
­Activity  199
Ventilation and Energy Demands During Physical Activity 290
Classification of Physical Activities by Energy Expenditure 199
Energy Cost of Breathing 295
The Met 200
Does Ventilation Limit Aerobic Power and Endurance
Daily Rates of Average Energy Expenditure 200
­Performance? 296
Energy Cost of Household, Industrial, and Recreational
Activities 202 PART 3 • ACID-BASE REGULATION 298
Heart Rate to Estimate Energy Expenditure 202 Buffering 298
Physiologic Buffers 300
Chapter 10 Effects of Intense Physical Activity 301
Energy Expenditure During Walking,
Chapter 15
Jogging, Running, and Swimming 205
The Cardiovascular System 303
Gross Versus Net Energy Expenditure 206
Economy of Human Movement 206 Cardiovascular System Components 304
Energy Expenditure During Walking 208 Hypertension 315
Energy Expenditure During Running 212 Blood Pressure Response to Physical Activity 318
Swimming 220 The Heart’s Blood Supply 320
Myocardial Metabolism 322
Chapter 11
Chapter 16
Individual Differences and Measurement
Cardiovascular Regulation and Integration 325
of Energy Capacities 227
Intrinsic Regulation of Heart Rate 326
Specificity Versus Generality of Metabolic Capacity and Exercise
Extrinsic Regulation of Heart Rate and Circulation 327
Performance 228
Distribution of Blood 333
Overview of Energy-Transfer Capacity During Exercise 228
Integrative Response During Physical Activity 336
Anaerobic Energy Transfer: The Immediate and Short-Term Energy
Physical Activity After Cardiac Transplantation 337
Systems 228
Aerobic Energy: The Long-Term Energy System 236 Chapter 17
Functional Capacity of the Cardiovascular
Section 3 Aerobic Systems System341
of Energy Delivery Cardiac Output 342
and Utilization 249 Cardiac Output at Rest 343
Cardiac Output During Physical Activity 344
Interview with Dr. Loring B. Rowell Cardiac Output Distribution 347
Chapter 12 Cardiac Output and Oxygen Transport 348
Pulmonary Structure and Function 253 Cardiovascular Adjustments to Upper-Body Exercise 352
Chapter 18
Surface Area and Gas Exchange 254
Anatomy of Ventilation 254 Skeletal Muscle: Structure and Function 355
Mechanics of Ventilation 255 Gross Structure of Skeletal Muscle 356
Lung Volumes and Capacities 258 Skeletal Muscle Ultrastructure 360
Lung Function, Aerobic Fitness, and Physical Performance 261 Muscle Fiber Alignment 362
Pulmonary Ventilation 261 Actin–Myosin Orientation 366
Variations From Normal Breathing Patterns 265 Chemical and Mechanical Events During Muscle Action
The Respiratory Tract During Cold-Weather Physical Activity 267 and Relaxation 367
Chapter 13 Muscle Fiber Type 374
Gas Exchange and Transport 269 Genes That Define Skeletal Muscle Phenotype 379
Fiber Type Differences Among Athletic Groups 379
PART 1 • GASEOUS EXCHANGE IN THE LUNGS
Chapter 19
AND TISSUES 270
Neural Control of Human Movement 383
Concentrations and Partial Pressures of Respired Gases 270
Gas Movement in Air and Fluids 271 Neuromotor System Organization 384
Gas Exchange in The Lungs and Tissues 272 Nerve Supply to Muscle 391
PART 2 • OXYGEN TRANSPORT 275 Motor Unit Functional Characteristics 396
Oxygen Transport in Blood 275 Receptors in Muscles, Joints, and Tendons: The Proprioceptors 400

McArdle9781451191554_FM.indd 12 12/11/2013 7:50:54 PM

 Contents xiii

Chapter 20 PART 1 • P
 harmacologic Agents for Ergogenic
The Endocrine System: Organization and Effects 548
Acute and Chronic Responses to Physical PART 2 • N
 onpharmacologic Approaches for
Activity407 Ergogenic ­Effects 571

Endocrine System Overview 408

Endocrine System Organization 408 Section 5 Exercise Performance
Resting and Exercise-Induced Endocrine Secretions 414 and Environmental
Gonadal Hormones 425 Stress593
Exercise Training and Endocrine Function 440
Resistance Training and Endocrine Function 445 Interview with Barbara Drinkwater
Opioid Peptides and Physical Activity 447 Chapter 24
Physical Activity, Infectious Illness, Cancer, and Immune
­Response 449
Physical Activity at Medium
and High Altitude 597
The Stress of Altitude 598
Part Two Acclimatization 600
Metabolic, Physiologic, and Exercise Capacities at Altitude 608
Applied Exercise Altitude Training and Sea-Level Performance 610
Physiology 455 Combine Altitude Stay With Low-Altitude Training 611
Chapter 25
Exercise and Thermal Stress 615
Section 4 Enhancement of Energy
Transfer Capacity 457 PART 1 • MECHANISMS OF THERMO­REGULATION 616
Thermal Balance 616
Interview with Bengt Saltin Hypothalamic Temperature Regulation 617
Chapter 21 Thermoregulation in Cold Stress: Heat Conservation
Training for Anaerobic and Aerobic Power 461 and Heat Production 617
Thermoregulation in Heat Stress: Heat Loss 618
Exercise Training Principles 462 Effects of Clothing on Thermoregulation 621
How Exercise Training Impacts the Anaerobic System 464 PART 2 • T
 HERMO­REGULATION AND ENVIRONMENTAL
Anaerobic System Changes with Training 464 HEAT STRESS DURING PHYSICAL ACTIVITY 626
How Training Impacts the Aerobic System 464 Physical Activity in The Heat 626
Factors That Affect Aerobic Training Responses 477 Maintaining Fluid Balance: Rehydration and Hyperhydration 630
How Long Before Improvements Occur? 484 Factors That Modify Heat Tolerance 632
Maintaining Gains in Aerobic Fitness 485 Complications From Excessive Heat Stress 635
Training Methods 486
PART 3 • T
 HERMO­REGULATION AND ENVIRONMENTAL
Overtraining: Too Much of a Good Thing 490
COLD STRESS DURING PHYSICAL ACTIVITY 637
Physical Activity During Pregnancy 491
Physical Activity in the Cold 637
Chapter 22 Cold Acclimatization 639
Muscular Strength: Training Muscles How Cold Is Too Cold? 640
to Become Stronger 499 Chapter 26
PART 1 • STRENGTH MEASUREMENT AND RESISTANCE Sport Diving 643
TRAINING 500 Diving History—Antiquity to the Present 644
Objectives of Resistance Training 502 Pressure–Volume Relationships and Diving Depth 652
Measurement of Muscle Strength 502 Snorkeling and Breath-Hold Diving 653
Gender Differences in Muscle Strength 506 Scuba Diving 659
Training Muscles to Become Stronger 509 Special Problems with Breathing Gases at High Pressures 661
PART 2 • STRUCTURAL AND FUNCTIONAL ADAPTATIONS Dives to Exceptional Depths: Mixed-Gas Diving 667
TO RESISTANCE TRAINING 528 Energy Cost of Underwater Swimming 670
Factors That Modify the Expression of Human
Strength 528 Chapter 27
Comparative Training Responses in Men and Women 535 Microgravity: The Last Frontier 673
Detraining Effects on Muscle 536 The Weightless Environment 674
Metabolic Stress of Resistance Training 537 Historical Overview of Aerospace Physiology and Medicine 680
Circuit Resistance Training 537 Modern Era 683
Muscle Soreness and Stiffness 538 Medical Evaluation for Astronaut Selection 686
Chapter 23 Bone 689
Special Aids to Exercise Training Countermeasure Strategies 702
and Performances 543 Overview of Physiologic Responses to Spaceflight 714
Nasa’s New Vision for the Future of Space Exploration 715
An Increasing Challenge To Fair Competition 544 Practical Benefits from Space Biology Research 720
On the Horizon 548 Final Words 722

McArdle9781451191554_FM.indd 13 12/11/2013 7:50:54 PM

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