H U M A N P H Y S I O L O G Y B R Y A N D E R R I C K S O N VALENCIA COLLEGE Director, Life Sciences Kevin Witt Executive Editor Bonnie Roesch Associate Editor Brittany Cheetham Developmental Editor Karen Trost Senior Marketing Manager Maria Guarascio Senior Product Designer Linda Muriello Senior Media Specialist Svetlana Barskaya Market Solutions Assistant Lindsey Myers Senior Production Editor Trish McFadden Photo Manager MaryAnne Price Illustration Coordinators Anna Melhorn/Claudia Durell Senior Designer Th omas Nery Th is book is printed on acid free paper. Founded in 1807, John Wiley & Sons, Inc. has been a valued source of knowledge and understanding for more than 200 years, helping people around the world meet their needs and fulfi ll their aspirations. Our company is built on a foundation of principles that include responsibility to the communities we serve and where we live and work. 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Upon completion of the review period, please return the evaluation copy to Wiley. Return instructions and a free of charge return shipping label are available at www.wiley.com/go/returnlabel. If you have chosen to adopt this textbook for use in your course, please accept this book as your complimentary desk copy. Outside of the United States, please contact your local sales representative. ISBN: 978-0-470-38140-3 Printed in the United States of America 10 9 8 7 6 5 4 3 2 1 Th e inside back cover will contain printing identifi cation and country of origin if omitted from this page. In addition, if the ISBN on the back cover diff ers from the ISBN on this page, the one on the back cover is correct. About the Author Bryan Derrickson is Professor of Biology at Valencia College in Orlando, Florida, where he teaches human anatomy and physi- ology as well as general biology and human sexuality. He received his bachelor’s degree in biology from Morehouse College and his Ph.D. in cell biology from Duke University. Bryan’s study at Duke was in the Physiology Division within the Department of Cell n Biology, so while his degree is in cell biology, his training focused o ks on physiology. At Valencia, he frequently serves on faculty hiring c erri committees. He has served as a member of the Faculty Senate, D n which is the governing body of the college, and as a member of a Bry the Faculty Academy Committee (now called the Teaching and of Learning Academy), which sets the standards for the acquisition y s e of tenure by faculty members. Nationally, he is a member of urt o the Human Anatomy and Physiology Society (HAPS) and the C National Association of Biology Teachers (NABT). Bryan has always wanted to teach. Inspired by several biology professors while in college, he decided to pursue physiology with an eye to teaching at the college level. He is completely dedicated to the success of his students. He particularly enjoys the challenges of his diverse student population, in terms of their age, ethnicity, and academic ability, and fi nds being able to reach all of them, despite their diff erences, a rewarding experience. His students continually recognize Bryan’s eff orts and care by nominating him for a campus award known as the “Valencia Professor Who Makes Valencia a Better Place to Start.” Bryan has received this award three times. In honor of my mother, Rosalind Gilmer Derrickson B.H.D. n o ks c erri D n a y Br of y es urt o C About the Author • iii Preface Welcome to your course in human physiology! Many of you are taking this course because you hope to pursue a career in medicine or one of the allied health professions. Others of you may be taking the course because you are simply interested in learning how your own amazing body functions. Whatever your motivations, Human Physiology, and WileyPLUS Learning Space have all the content and tools needed to ensure that you receive a solid foundation and the knowledge and skills to reach your desired goals. I am passionate about the discipline of physiology, and have an equally strong passion for teaching. Physiology is a fascinating subject and I enjoy conveying this information to students and guiding them through the intricacies of the many complex functions of the human body. I wrote this text because of my desire to share the story of physiology with a wider audience. Human Physiology is a comprehensive text that uses four main underlying principles in physiology as a foundation for specifi c details of all the systems of the human body. Th ese principles include home- ostasis, mechanisms of action, communication, and integration. As you progress through the text, you will discover these underlying themes supporting your understanding of core physiological concepts. Most importantly, I endeavored to distinguish this text by uniquely combining three powerful ele- ments: (1) clear, easy-to-follow writing style supported with carefully developed fi gures; (2) an emphasis on the development of vital critical thinking skills; and (3) a fully integrated digital platform rich in interactive activities and media. Together these elements provide a superior level of coverage that helps prepare you for successful careers in medicine and allied health. THE NARRATIVE AND VISUALS Each chapter in Human Physiology is written in a style that is very straightforward and easy to under- stand. Th is type of writing style facilitates comprehension and retention of key physiological concepts. Words can convey a lot when carefully chosen, but if you are like most students today, visual represen- tation of the material is of equal importance to you. You’ll fi nd clear visual explanations that bring the words on the page to life. In addition, each chapter is fi lled with boxed information on relevant clinical connections and other real-life examples that will stir your interest in, and solidify understanding of, the relevant science at hand. CRITICAL THINKING Understanding how to think critically about the scientifi c information presented is vital to your success. Human Physiology, 1st edition and WileyPLUS Learning Space have several features that give you the opportunity to hone this essential skill. Critical Th inking exercises help you to think logically and critical- ly about real-life scenarios that involve important physiological concepts. Research to Reality is a unique feature that provides you with the opportunity to analyze and interpret real scientifi c data from primary research articles. Physiological Equation boxes describe key physiological equations that you can use to analyze and quantify certain physiological parameters. Ponder Th is questions at the end of each chapter provide additional opportunities to think critically about physiological information. ENGAGING DIGITALLY I am so excited to have my text fully integrated with WileyPLUS Learning Space. Th is platform allows you to create a personalized study plan, assess your progress along the way, and make deeper con- nections as you interact with the course material and with one another. Th is collaborative learning environment provides immediate insight into both your strengths and your problem areas through a combination of dynamic course materials—such as 3D animations, game-like exercises, and laboratory simulations—and visual reports so you can act on what’s most important to help you master the course material. WileyPLUS Learning Space also includes ORION—integrated, adaptive practice that helps you build profi ciency on particular topics and use your study time most eff ectively. iv • Preface Includes ORION Adaptive Practice An easy way to help students learn, collaborate, and grow. Designed to engage today’s student, WileyPLUS Learning Space will transform any course into a vibrant, collaborative, learning community. Photo credit: Sam Edwards/Getty Images Identify which students FFaacilitate student Measure outcomes are struggling early in the ennggagement both in and to promote continuous semester. ouuttside of class. improvement. Educators assess the real-time Educaators can quickly organize With visual reports, it’s easy for engagement and performance of learninnngg activities, manage student both students and educators to each student to inform teaching collaborraaation, and customize gauge problem areas and act on decisions. Students always know their courrsssee. what’s most important. what they need to work on. www.wileypluslearningspace.com Acknowledgments Human Physiology, 1st edition and WileyPLUS Learning Space would Space with ORION. Th eir thoughtful execution of resources and as- not be possible without the help of many, particularly the numerous sessment has raised the bar for excellence. Th ank you, so very much. academic colleagues that collaborated with us along the way. First to Brian Antonsen Marshall University thank are three contributors whose work informed and enhanced our Heidi Bustamante University of Colorado, Boulder focus on critical thinking and connections to real life situations and Patrick Caff erty Emory University activities: Janet Casagrand University of Colorado, Boulder David Mallory, Marshall University, was instrumental in the design Lynn Diener Mount Mary University and execution of the Critical Th inking Boxes and step-by-step ques- tions found in each chapter. His choice of topics for these boxes John Erickson Ivy Tech State College-SE adds great interest and really highlights the connections to the Gregory Loftin Metropolitan Community College concepts at hand. Th ank you so much, David. of Kansas City Lynn Diener, Mount Mary University, researched and chose most Geoff Goellner Minnesota State University of the primary research papers used in the Research to Reality Michael Griffi n Angelo State University activities. She used her expertise and experience in implementing Jill Tall Youngstown State University this kind of investigation in her own classes to bring this unique Tracy Wagner Washburn University of Topeka feature to this project. I am very grateful, Lynn. Paul Wagner Washburn University of Topeka Heidi Bustamante, University of Colorado, Boulder, excels at writ- Chad Wayne University of Houston ing thoughtful questions that challenge students to critically think about what they have learned. I am so happy that she chose to en- Th e development of a fi rst edition text and media for a course as hance this text by writing the end-of-chapter Ponder Th is questions. complex as human physiology is a long process and would not be pos- I really appreciate your work, Heidi. sible without the continued involvement of those “in the trenches” I am also very grateful that Wiley commissioned a board of aca- teaching the course, who guided and informed our choices all along demic advisors with expertise in teaching physiology to help inform the way. I am very grateful to my colleagues who have reviewed the not only individual resources integrated with this text, but also as a drafts of manuscript, participated in focus groups and workshops, or sounding board for all of us as we progressed through development. off ered suggestions for improvement. Ateegh Al-Arabi Johnson County Community College WILEY ADVISORY BOARD FOR ANATOMY Brenda Alston-Mills Michigan State University AND PHYSIOLOGY Gwen Bachman University of Nebraska, Lincoln Heidi Bustamante University of Colorado, Boulder Ari Berowitz University of Oklahoma, Norman Patrick Caff erty Emory University Eric Blough Marshall University Janet Casagrand University of Colorado, Boulder Sunny K. Boyd University of Notre Dame Lynn Diener Mount Mary University Carol Britson University of Mississippi John Erickson Ivy Tech State College-SE Jackie Brittingham Simpson College Melaney Farr Salt Lake Community College Brent Bruot Kent State University Geoff Goellner Minnesota State University Heidi Bustamante University of Colorado, Boulder Michael Griffi n Angelo State University Sherell Byrd Fort Lewis College Wanda Hargroder Louisiana State University Martin Burg Grand Valley State University DJ Hennager Kirkwood Community College Phyllis Callahan Miami University Sandra Hutchinson Santa Monica College Jackie Carnegie University of Ottawa Heather Labbe University of Montana - Missoula Robert Carroll East Carolina University Tom Lancraft St. Petersburg College Janet Casagrand University of Colorado, Boulder David Mallory Marshall University Daniel Castellanos Florida International University Russell Nolan Baton Rouge Community College Chris Dewitt University of South Carolina Aiken Terry Th ompson Wor-Wic Community College Debora Christensen Drake University Tracy Wagner Washburn University of Topeka Ruth Clark Washington University, St. Louis Paul Wagner Washburn University of Topeka Josefa Cubina New York Institute of Technology I thank the following group of academics who have contributed Maria de Bellard California State University, Northridge to the creation and integration of this text with WileyPLUS Learning Michael Deschenes College of William and Mary vi • Acknowledgments Lynn Diener Mount Mary University David Nutting University of Tennessee, Memphis Stephen Dodd University of Florida Linda Ogren University of California - Santa Cruz John Erickson Ivy Tech Ok-Kyong Park-Sarge University of Kentucky Max G. Ervin Middle Tennessee State University William Percy University of South Dakota Carol Fassbinder-Orth Creighton University David Pistole Indiana University of Pennsylvania Ralph Ferges Palomar College Robert Preston Illinois State University James Ferraro Southern Illinois School of Medicine David Petzel Creighton University Michael Finkler Indiana University - Kokomo Steve Price Virginia Commonwealth University John Fishback Ozarks Tech Community College Peter Reiser Ohio State University David Flory University of Central Florida Nick A. Ritucci Wright State University Victor Fomin University of Delaware Laurel Roberts University of Pittsburgh Michelle French University of Toronto Sonia Rocah-Sanchez Creighton University Geoff Goellner Minnesota State University Dean Scherer Oklahoma State University Barbara Goodman University of South Dakota David Schulz University of Missouri Mike Griffi n Angelo State University Virginia Shea University of North Carolina, Chapel Hill Bryan Hamilton Waynesburg University Allison Shearer Grossmont College Janet Haynes Long Island University Rachel Smetanka Southern Utah University Susan Heaphy Th e Ohio State University at Lima James Strauss Pennsylvania State University Steven Heidemann Michigan State University Erik Swenson University of Washington Stephen Henderson California State University, Chico Jill Tall Youngstown State University William Higgins University of Maryland Bonnie Tarricone Ivy Tech Community College at Indianapolis Steven Hobbs University of Colorado Mark Th omas University of Northern Colorado James Hoff man Diablo Valley College Maureen Tubbiola St. Cloud State University Kelly Johnson University of Kansas Paul Wagner Washburn University of Topeka Steven King Oregon Health & Science University Tracy Wagner Washburn University of Topeka Jennifer Kneafsey Tulsa Community College - Curt Walker Dixie State College of Utah Northeast Campus Richard Walker University of Calgary Megan Knoch Indiana University of Pennsylvania R. Douglass Watson University of Alabama, Birmingham Joan Lafuze Indiana University East Chad Wayne University of Houston Dean Lauritzen City College of San Francisco Last, but certainly not least, I owe tremendous gratitude to the John Lepri University of North Carolina, Greensboro team at Wiley. Th is team of collaborative publishing professionals is Paul Lonquich California State University, Northridge dedicated, enthusiastic, and talented and each brings a level of skill to Jennifer Lundmark California State University - Sacramento the job that, I believe, is unparalleled. My hat is off to the entire team: David Mallory Marshall University Bonnie Roesch, Executive Editor; Karen Trost, Developmental Editor; Brittany Cheetham, Associate Editor; Maria Guarascio, Senior Market- Th eresa Martin San Mateo Community College ing Manager; Linda Muriello, Senior Product Designer; Lindsey Myers, Michael Masson Santa Barbara Community College Market Solutions Assistant; Trish McFadden, Senior Production Tamara Mau University of California, Berkeley Editor; Mary Ann Price, Senior Photo Editor; Claudia Volano and Anna Eric McElroy College of Charleston Melhorn, Illustration Coordinators; Th omas Nery, Senior Designer. I Th omas McNeilis Dixie State College of Utah also want to acknowledge Kevin Witt, Director, for his support of this John McReynolds University of Michigan project. My heartfelt thanks to you all! Jamie Melling Western University Bryan Derrickson Jeanne Mitchell Truman State University Department of Science, PO Box 3028 Diane Morel University of the Sciences Valencia College Orlando, FL 32802 Tim Mullican Dakota Wesleyan University [email protected] Cheryl Neudauer Minneapolis Community and Technical College Acknowledgments • vii Brief Contents 1 An Introduction to Physiology 1 2 Chemical Composition of the Body 19 3 Cells 56 4 Metabolism 95 5 Transport Across the Plasma Membrane 128 6 Cell Signaling 156 7 The Nervous System and Neuronal Excitability 187 8 The Central Nervous System 238 9 Sensory Systems 275 10 Autonomic and Somatic Nervous Systems 341 11 Muscle 364 12 Control of Body Movement 411 13 The Endocrine System 432 14 The Cardiovascular System: The Heart 484 15 The Cardiovascular System: Blood Vessels and Hemodynamics 524 16 The Cardiovascular System: The Blood 565 17 The Immune System 593 18 The Respiratory System 625 19 The Urinary System 664 20 Fluid, Electrolyte, and Acid–Base Homeostasis 702 21 The Digestive System 721 22 Metabolic Adaptation, Energy Balance, and Temperature Regulation 764 23 The Reproductive Systems 781 Appendix A: Measurements A-1 Appendix B: Periodic Table B-3 Appendix C: Normal Values for Selected Blood Tests C-4 Appendix D: Normal Values for Selected Urine Tests D-6 Glossary G-1 Index I-1 viii • Brief Contents