,FOURTEENTH EDITION VA N D E R ’ S
Human
Physiology
The Mechanisms of Body Function
ERIC P. WIDMAIER
B OS TO N U N I V E R S I T Y
HERSHEL RAFF
M E D I C A L CO L L E G E O F W I S CO N S I N
AU R O R A S T. LU K E ’ S M E D I C A L C E N T E R
KEVIN T. STRANG
U N I V E R S I T Y O F W I S CO N S I N – M A D I S O N
,VANDER’S HUMAN PHYSIOLOGY: THE MECHANISMS OF BODY FUNCTION, FOURTEENTH EDITION
Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2016 by McGraw-Hill Education.
All rights reserved. Printed in the United States of America. Previous editions © 2014, 2011, and 2008. No part of this
publication may be reproduced or distributed in any form or by any means, or stored in a database or retrieval system, without
the prior written consent of McGraw-Hill Education, including, but not limited to, in any network or other electronic storage or
transmission, or broadcast for distance learning.
Some ancillaries, including electronic and print components, may not be available to customers outside the United States.
This book is printed on acid-free paper.
1 2 3 4 5 6 7 8 9 0 DOW/DOW 1 0 9 8 7 6 5
ISBN 978–1–259–29409–9
MHID 1–259–29409–9
Senior Vice President, Products & Markets: Kurt L. Strand Content Licensing Specialists: Lori Hancock/Lorraine Buczek
Vice President, General Manager, Products & Cover Image: © Roxana Wegner/Getty Images (X-Ray head
Markets: Marty Lange scan); © Hero Images/Getty Images (Doctor and Patient);
Vice President, Content Design & © Science Photo Library–Steve Gschmeissner/Getty
Delivery: Kimberly Meriwether David Images (Nerve fiber); © Science Photo Library–Steve
Managing Director: Michael S. Hackett Gschmeissner/Getty Images (Colored SEM); © Clouds
Brand Manager: Amy Reed/Chloe Bouxsein Hill Imaging LTD/Science Photo Library/Getty Images
Director, Product Development: Rose Koos (Human heart and lung blood vessels); © skynesher/Getty
Product Developer: Fran Simon Images (Group of runners); © Science Photo Library–
Marketing Manager: James Connely Steve Gschmeissner/Getty Images (Blood Clot SEM);
Director, Content Design & Delivery: Linda Avenarius © Comotion Design/Getty Images (X-Ray human heart
Program Manager: Angela R. Fitz Patrick and vascular system)
Content Project Managers: Vicki Krug/Brent dela Cruz Compositor: SPi Global
Buyer: Sandy Ludovissy Printer: R. R. Donnelley
Design: Matt Backhaus
All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.
NOTICE
Medicine is an ever-changing science. As new research and clinical experience broaden our knowledge, changes in treatment
and drug therapy are required. The authors and the publisher of this work have checked with sources believed to be reliable
in their efforts to provide information that is complete and generally in accord with the standards accepted at the time of
publication. However, in view of the possibility of human error or changes in medical sciences, neither the authors nor the
publisher nor any other party who has been involved in the preparation or publication of this work warrants that the information
contained herein is in every respect accurate or complete, and they disclaim all responsibility for any errors or omissions or for
the results obtained, from use of the information contained in this work. Readers are encouraged to confirm the information
contained herein with other sources. For example and in particular, readers are advised to check the product information
sheet included in the package of each drug they plan to administer to be certain that the information contained in this work is
accurate and that changes have not been made in the recommended dose or in the contraindications for administration. This
recommendation is of particular importance in connection with new or infrequently used drugs.
Library of Congress Cataloging-in-Publication Data
Names: Widmaier, Eric P. | Raff, Hershel, 1953- | Strang, Kevin T. | Vander,
Arthur J., 1933- Human physiology.
Title: Human physiology : the mechanisms of body function / Eric P. Widmaier,
Boston University, Hershel Raff, Medical College of Wisconsin, Aurora St.
Luke’s Medical Center, Kevin T. Strang, University of Wisconsin-Madison.
Other titles: Human physiology
Description: Fourteenth edition. | New York, NY : McGraw-Hill, [2016] |
Includes bibliographical references and index.
Identifiers: LCCN 2015024583 | ISBN 9781259294099 (alk. paper)
Subjects: LCSH: Human physiology.
Classification: LCC QP34.5 .W47 2016 | DDC 612—dc23
LC record available at http://lccn.loc.gov/2015024583
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate
an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education does not guarantee the accuracy of the
information presented at these sites.
mheducation.com/highered
, Meet the Authors
ERIC P. WIDMAIER received his Ph.D. in 1984 in Endocrinology from the University of
California at San Francisco. His postdoctoral training was in endocrinology and physiology at the
Worcester Foundation for Experimental Biology in Shrewsbury, Massachusetts, and The Salk Institute
in La Jolla, California. His research is focused on the control of body mass and metabolism in mammals,
the mechanisms of hormone action, and molecular mechanisms of intestinal and hypothalamic
adaptation to high-fat diets. He is currently Professor of Biology at Boston University, where he teaches
Human Physiology and has been recognized with the Gitner Award for Distinguished Teaching by the
College of Arts and Sciences, and the Metcalf Prize for Excellence in Teaching by Boston University. He
is the author of many scientific and lay publications, including books about physiology for the general
reader. He lives outside Boston with his wife Maria and children Caroline and Richard.
H ER SH EL R AFF received his Ph.D. in Environmental Physiology from the Johns Hopkins
University in 1981 and did postdoctoral training in Endocrinology at the University of California
at San Francisco. He is now a Professor of Medicine (Endocrinology, Metabolism, and Clinical
Nutrition), Surgery, and Physiology at the Medical College of Wisconsin and Director of the
Endocrine Research Laboratory at Aurora St. Luke’s Medical Center. He teaches physiology to
medical and graduate students as well as clinical fellows. At the Medical College of Wisconsin,
he is the Endocrinology/Reproduction Unit Director for the new integrated curriculum. He was
an inaugural inductee into the Society of Teaching Scholars, received the Beckman Basic Science
Teaching Award three times, received the Outstanding Teacher Award from the Graduate School,
and has been one of the MCW’s Outstanding Medical Student Teachers in multiple years. He is also
an Adjunct Professor of Biomedical Sciences at Marquette University. Dr. Raff’s basic research
focuses on the adaptation to low oxygen (hypoxia). His clinical interest focuses on pituitary and
adrenal diseases, with a special focus on laboratory tests for the diagnosis of Cushing’s syndrome.
He resides outside Milwaukee with his wife Judy and son Jonathan.
K EV I N T. STR ANG received his Master’s Degree in Zoology (1988) and his Ph.D. in
Physiology (1994) from the University of Wisconsin at Madison. His research area is cellular
mechanisms of contractility modulation in cardiac muscle. He teaches a large undergraduate
systems physiology course as well as first-year medical physiology in the UW-Madison School of
Medicine and Public Health. He was elected to UW-Madison’s Teaching Academy and as a Fellow
of the Wisconsin Initiative for Science Literacy. He is a frequent guest speaker at colleges and high
schools on the physiology of alcohol consumption. He has twice been awarded the UW Medical
Alumni Association’s Distinguished Teaching Award for Basic Sciences, and also received the
University of Wisconsin System’s Underkofler/Alliant Energy Excellence in Teaching Award. In
2012 he was featured in The Princeton Review publication, “The Best 300 Professors.” Interested
in teaching technology, Dr. Strang has produced numerous animations of figures from Vander’s
Human Physiology available to instructors and students. He lives in Madison with his wife Sheryl
and his children Jake and Amy.
T O O U R FA M I L I E S : M A R I A , C A R O L I N E , A N D R I C H A R D ; J U DY A N D J O N A T H A N ;
S H E RY L , J A K E , A N D A M Y
iii
Human
Physiology
The Mechanisms of Body Function
ERIC P. WIDMAIER
B OS TO N U N I V E R S I T Y
HERSHEL RAFF
M E D I C A L CO L L E G E O F W I S CO N S I N
AU R O R A S T. LU K E ’ S M E D I C A L C E N T E R
KEVIN T. STRANG
U N I V E R S I T Y O F W I S CO N S I N – M A D I S O N
,VANDER’S HUMAN PHYSIOLOGY: THE MECHANISMS OF BODY FUNCTION, FOURTEENTH EDITION
Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2016 by McGraw-Hill Education.
All rights reserved. Printed in the United States of America. Previous editions © 2014, 2011, and 2008. No part of this
publication may be reproduced or distributed in any form or by any means, or stored in a database or retrieval system, without
the prior written consent of McGraw-Hill Education, including, but not limited to, in any network or other electronic storage or
transmission, or broadcast for distance learning.
Some ancillaries, including electronic and print components, may not be available to customers outside the United States.
This book is printed on acid-free paper.
1 2 3 4 5 6 7 8 9 0 DOW/DOW 1 0 9 8 7 6 5
ISBN 978–1–259–29409–9
MHID 1–259–29409–9
Senior Vice President, Products & Markets: Kurt L. Strand Content Licensing Specialists: Lori Hancock/Lorraine Buczek
Vice President, General Manager, Products & Cover Image: © Roxana Wegner/Getty Images (X-Ray head
Markets: Marty Lange scan); © Hero Images/Getty Images (Doctor and Patient);
Vice President, Content Design & © Science Photo Library–Steve Gschmeissner/Getty
Delivery: Kimberly Meriwether David Images (Nerve fiber); © Science Photo Library–Steve
Managing Director: Michael S. Hackett Gschmeissner/Getty Images (Colored SEM); © Clouds
Brand Manager: Amy Reed/Chloe Bouxsein Hill Imaging LTD/Science Photo Library/Getty Images
Director, Product Development: Rose Koos (Human heart and lung blood vessels); © skynesher/Getty
Product Developer: Fran Simon Images (Group of runners); © Science Photo Library–
Marketing Manager: James Connely Steve Gschmeissner/Getty Images (Blood Clot SEM);
Director, Content Design & Delivery: Linda Avenarius © Comotion Design/Getty Images (X-Ray human heart
Program Manager: Angela R. Fitz Patrick and vascular system)
Content Project Managers: Vicki Krug/Brent dela Cruz Compositor: SPi Global
Buyer: Sandy Ludovissy Printer: R. R. Donnelley
Design: Matt Backhaus
All credits appearing on page or at the end of the book are considered to be an extension of the copyright page.
NOTICE
Medicine is an ever-changing science. As new research and clinical experience broaden our knowledge, changes in treatment
and drug therapy are required. The authors and the publisher of this work have checked with sources believed to be reliable
in their efforts to provide information that is complete and generally in accord with the standards accepted at the time of
publication. However, in view of the possibility of human error or changes in medical sciences, neither the authors nor the
publisher nor any other party who has been involved in the preparation or publication of this work warrants that the information
contained herein is in every respect accurate or complete, and they disclaim all responsibility for any errors or omissions or for
the results obtained, from use of the information contained in this work. Readers are encouraged to confirm the information
contained herein with other sources. For example and in particular, readers are advised to check the product information
sheet included in the package of each drug they plan to administer to be certain that the information contained in this work is
accurate and that changes have not been made in the recommended dose or in the contraindications for administration. This
recommendation is of particular importance in connection with new or infrequently used drugs.
Library of Congress Cataloging-in-Publication Data
Names: Widmaier, Eric P. | Raff, Hershel, 1953- | Strang, Kevin T. | Vander,
Arthur J., 1933- Human physiology.
Title: Human physiology : the mechanisms of body function / Eric P. Widmaier,
Boston University, Hershel Raff, Medical College of Wisconsin, Aurora St.
Luke’s Medical Center, Kevin T. Strang, University of Wisconsin-Madison.
Other titles: Human physiology
Description: Fourteenth edition. | New York, NY : McGraw-Hill, [2016] |
Includes bibliographical references and index.
Identifiers: LCCN 2015024583 | ISBN 9781259294099 (alk. paper)
Subjects: LCSH: Human physiology.
Classification: LCC QP34.5 .W47 2016 | DDC 612—dc23
LC record available at http://lccn.loc.gov/2015024583
The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate
an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education does not guarantee the accuracy of the
information presented at these sites.
mheducation.com/highered
, Meet the Authors
ERIC P. WIDMAIER received his Ph.D. in 1984 in Endocrinology from the University of
California at San Francisco. His postdoctoral training was in endocrinology and physiology at the
Worcester Foundation for Experimental Biology in Shrewsbury, Massachusetts, and The Salk Institute
in La Jolla, California. His research is focused on the control of body mass and metabolism in mammals,
the mechanisms of hormone action, and molecular mechanisms of intestinal and hypothalamic
adaptation to high-fat diets. He is currently Professor of Biology at Boston University, where he teaches
Human Physiology and has been recognized with the Gitner Award for Distinguished Teaching by the
College of Arts and Sciences, and the Metcalf Prize for Excellence in Teaching by Boston University. He
is the author of many scientific and lay publications, including books about physiology for the general
reader. He lives outside Boston with his wife Maria and children Caroline and Richard.
H ER SH EL R AFF received his Ph.D. in Environmental Physiology from the Johns Hopkins
University in 1981 and did postdoctoral training in Endocrinology at the University of California
at San Francisco. He is now a Professor of Medicine (Endocrinology, Metabolism, and Clinical
Nutrition), Surgery, and Physiology at the Medical College of Wisconsin and Director of the
Endocrine Research Laboratory at Aurora St. Luke’s Medical Center. He teaches physiology to
medical and graduate students as well as clinical fellows. At the Medical College of Wisconsin,
he is the Endocrinology/Reproduction Unit Director for the new integrated curriculum. He was
an inaugural inductee into the Society of Teaching Scholars, received the Beckman Basic Science
Teaching Award three times, received the Outstanding Teacher Award from the Graduate School,
and has been one of the MCW’s Outstanding Medical Student Teachers in multiple years. He is also
an Adjunct Professor of Biomedical Sciences at Marquette University. Dr. Raff’s basic research
focuses on the adaptation to low oxygen (hypoxia). His clinical interest focuses on pituitary and
adrenal diseases, with a special focus on laboratory tests for the diagnosis of Cushing’s syndrome.
He resides outside Milwaukee with his wife Judy and son Jonathan.
K EV I N T. STR ANG received his Master’s Degree in Zoology (1988) and his Ph.D. in
Physiology (1994) from the University of Wisconsin at Madison. His research area is cellular
mechanisms of contractility modulation in cardiac muscle. He teaches a large undergraduate
systems physiology course as well as first-year medical physiology in the UW-Madison School of
Medicine and Public Health. He was elected to UW-Madison’s Teaching Academy and as a Fellow
of the Wisconsin Initiative for Science Literacy. He is a frequent guest speaker at colleges and high
schools on the physiology of alcohol consumption. He has twice been awarded the UW Medical
Alumni Association’s Distinguished Teaching Award for Basic Sciences, and also received the
University of Wisconsin System’s Underkofler/Alliant Energy Excellence in Teaching Award. In
2012 he was featured in The Princeton Review publication, “The Best 300 Professors.” Interested
in teaching technology, Dr. Strang has produced numerous animations of figures from Vander’s
Human Physiology available to instructors and students. He lives in Madison with his wife Sheryl
and his children Jake and Amy.
T O O U R FA M I L I E S : M A R I A , C A R O L I N E , A N D R I C H A R D ; J U DY A N D J O N A T H A N ;
S H E RY L , J A K E , A N D A M Y
iii
