ir YOU’VE JUST PURCHASED MORE THAN A TEXTBOOK! Evolve Student Resources for Klein: Cunningham’s Textbook of Veterinary Physiology, Sixth Edition, include the following: • Animations • Clinical Correlations • Glossary • Practice Tests Activate the complete learning experience that comes with each NEW textbook purchase by registering with your scratch-off access code at http://evolve.com/Klein/physiology/ If you rented or purchased a used book and the scratch-off code at right has already been revealed, the code may have been used and cannot be re-used for registration. To purchase a new code to access these valuable study resources, simply follow the link above. Place Sticker Here REGISTER TODAY! You can now purchase Elsevier products on Evolve! Go to evolve.com/shop to search and browse for products.ir Cunningham’s Textbook of VETERINARY PHYSIOLOGY Sixth Edition Bradley G. Klein, PhD Associate Professor of Neuroscience Department of Biomedical Sciences and Pathobiology Virginia-Maryland College of Veterinary Medicine Virginia Polytechnic Institute and State University Blacksburg, Virginia 3251 Riverport Lane VetBooks.
Louis, Missouri 63043 CUNNINGHAM’S TEXTBOOK OF VETERINARY PHYSIOLOGY, ISBN: 978-0-323-55227-1 SIXTH EDITION Copyright © 2020 by Elsevier, Inc. All rights reserved. Previous edition copyrighted 2013, 2007, 2002, 1997, 1992. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher.
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To the fullest extent of the law, no responsibility is assumed by Elsevier, authors, editors or contributors for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions, or ideas contained in the material herein. ISBN 978-0-323-55227-1 Senior Content Strategist: Jennifer Flynn-Briggs Senior Content Development Manager: Ellen Wurm-Cutter Content Development Specialist: Kathleen Nahm Publishing Services Manager: Deepthi Unni Senior Project Manager: Kamatchi Madhavan Project Manager: Haritha Dharmarajan Designer: Bridget Hoette Printed in China Last digit is the print number 9 8 7 6 5 4 3 2 1 VetBooks.ir In loving and lasting memory of Ira Matthew Klein (1985-2013). Ansar Ahmed, DVM, PhD Bradley G. Klein, PhD Professor and Associate Dean for Research Associate Professor of Neuroscience Department of Biomedical Sciences and Pathobiology Department of Biomedical Sciences and Pathobiology Virginia-Maryland College of Veterinary Medicine Virginia-Maryland College of Veterinary Medicine Virginia Polytechnic Institute and State University Virginia Polytechnic Institute and State University Blacksburg, Virginia Blacksburg, Virginia Steven P.
Brinsko, DVM, MS, PhD, DACT Xin M. Luo, MS, PhD Professor and Associate Department Head of Academics Associate Professor of Immunology Department of Large Animal Clinical Sciences Department of Biomedical Sciences and Pathobiology College of Veterinary Medicine and Biomedical Sciences Virginia-Maryland College of Veterinary Medicine Texas A&M University Virginia Polytechnic Institute and State University College Station, Texas Blacksburg, Virginia Autumn P. Davidson, DVM, MS, DACVIM (Internal Brian K. Petroff, DVM, PhD Medicine) Associate Professor and Section Chief, Endocrinology Clinical Professor Department of Pathobiology and Diagnostic Investigation Veterinary Medical Teaching Hospital Diagnostic Center for Population and Animal Health Department of Medicine and Epidemiology College of Veterinary Medicine School of Veterinary Medicine Michigan State University University of California-Davis East Lansing, Michigan Davis, California Juan E.
Romano, DVM, MS, PhD, DACT Susan L. Ewart, DVM, PhD, DACVIM Professor Professor and Special Assistant to the Dean Department of Large Animal Clinical Sciences Department of Large Animal Clinical Sciences College of Veterinary Medicine and Biomedical Sciences College of Veterinary Medicine Texas A&M University Michigan State University College Station, Texas East Lansing, Michigan John H., DVM, MS, DACVIM Deborah S. Greco, DVM, PhD, DACVIM-SAIM Professor Senior Research Scientist Neurology and Neurosurgery Nestle Purina Petcare Department of Small Animal Clinical Sciences St. Louis, Missouri Virginia-Maryland College of Veterinary Medicine Virginia Polytechnic Institute and State University Steven R.
Heidemann, PhD Blacksburg, Virginia Professor Emeritus Department of Physiology Gerhardt G. Schurig, DVM, MS, PhD Michigan State University Professor and International Programs Strategist East Lansing, Michigan Department of Biomedical Sciences and Pathobiology Virginia-Maryland College of Veterinary Medicine Thomas H. Herdt, DVM, MS, DACVIM, DACVN Virginia Polytechnic Institute and State University Professor and Chief of Nutrition Blacksburg, Virginia Department of Large Animal Clinical Sciences and Veterinary Diagnostic Laboratory College of Veterinary Medicine Michigan State University East Lansing, Michigan iv Contributors v George H. Stabenfeldt, DVM, PhD (deceased) Jill W.
Verlander, DVM Professor Scientist Department of Reproduction Department of Medicine VetBooks.ir School of Veterinary Medicine Division of Nephrology, Hypertension, and Renal University of California-Davis Transplantation Davis, California University of Florida College of Medicine (Contributor of past editions) Gainesville, Florida Robert B. Stephenson, PhD Sharon G. Witonsky, DVM, PhD, DACVIM Associate Professor Associate Professor Department of Physiology Equine Field Service Michigan State University Department of Large Animal Clinical Sciences East Lansing, Michigan Virginia-Maryland College of Veterinary Medicine Virginia Polytechnic Institute and State University Blacksburg, Virginia Preface VetBooks.ir Physiology is the study of the normal functions of the body—the Instructors will appreciate the items in the illustration bank, which study of the body’s molecules, cells, and organ systems, and the can be downloaded for lecture use. There is also a Glossary covering interrelationships among them.
Because the study of medicine is many of the italicized words from the printed text. the study of abnormal functions of the body, it is essential to Immense gratitude is due to the contributing authors who have understand normal physiology if one is to understand the mecha- worked hard to ensure that the information in this latest edition nisms of disease. For this reason, physiology and other important is accurate and up-to-date. This has resulted in new illustrations sciences basic to medicine are introduced early in the veterinary and Clinical Correlations, as well as addition of material on new curriculum., a brain lymphatic system; neutrophil extracellular Physiology is a vast subject, and the normal time constraints traps).
The expertise of two additional section authors is welcomed, on veterinary students are usually prohibitive for learning all that Dr. Ewart for Respiratory Function and Homeostasis, is known about it. Therefore an effort was made to limit the and Dr. Petroff for Endocrinology.
Luo concepts presented in this book to those germane to the practice contributed some material to the Immunology section. Notable of veterinary medicine. Because the scope of physiology encompasses recognition is in order for Dr. Edward Robinson, who is retiring many scientific disciplines and levels of analysis, the contributing from this project after contributing to all of its previous editions, authors represent not only the field of physiology but others, among and who authored the initial material on respiratory physiology them neuroscience, cell biology, molecular biology, immunology, and on homeostasis.
Thanks also to Dr. Ayman Sayegh for contribut- and various clinical specialties. Most of these authors are also ing to an earlier edition of the introductory chapter on regulation veterinarians or have consulted with veterinary clinicians regarding of gastrointestinal function. Suggestions of ways to improve this content.
Sections on the immune system and cancer underscore text in subsequent editions are always welcome. the intimate relationship between the understanding of cell and Particular gratitude is due to the book’s medical illustrator, molecular biology, physiological function, and veterinary Jeanne Robertson, who drew the new illustrations for this edition medicine. and revised much of the preexisting artwork. Thanks are also in This book is designed for first-year veterinary students.
The order for the folks at Elsevier who were instrumental in producing goal is to introduce the student to the principles and concepts of the sixth edition, among them Penny Rudolph, Jennifer Flynn- physiology pertinent to the practice of veterinary medicine. Other Briggs, Ellen Wurm-Cutter, Anna Miller, Madhavan Kamatchi, goals are to introduce the reader to physiopathology and clinical and Haritha Dharmarajan. Special recognition is due for Kathleen problem-solving techniques and to help the reader understand the Nahm, the Elsevier content development specialist who shepherded relationship between physiology and the practice of veterinary almost all aspects of this project through various trials and tribula- medicine. tions and who did an admirable job managing my not infrequent This book is designed to be as student friendly as possible.
New tardiness yet otherwise fastidious nature. Jonathan Abbott, concepts in the text are introduced by a declarative statement Virginia Buechner-Maxwell, and Shireen Hafez unselfishly provided designed to summarize the essential point. This format also helps their valuable opinions on various aspects of the book that resulted the reader survey the chapter or review for an examination. These in its improvement.
Bonnie Smith was particularly helpful in declarative statements are also listed at the beginning of the chapter providing advice on the accuracy of the cover illustration. Again, as an outline of Key Points. this book would not exist without the invaluable expertise of the Chapters include Clinical Correlations at the end. Concordant section authors who worked so hard to make this the best veterinary with the increasing trend toward early integration of basic science physiology text possible.
and clinical material in veterinary curricula, these Clinical Correla- The most immense debt is due to Dr. Jim Cunningham, whose tions are designed to show the reader how knowledge of physiology vision, guidance, and expertise made the Textbook of Veterinary is applied to the diagnosis and treatment of veterinary patients. Physiology a reality and a success, and has made “Cunningham’s” They also provide the student with an additional way to think a household name among veterinary students and faculty of several through the principles and concepts presented, and they can serve generations. Placing his name in the book’s title is in partial recogni- as a basis for classroom case discussions.
tion of that debt. He is also recognized here for authoring the Several Practice Questions are included in each chapter as another initial material on neurophysiology upon which I have built over method for students to review the book’s content. The Bibliography the years. The instructional style he instituted continues in this for each chapter is designed to lead the reader to more advanced edition and will continue in future editions of the text.
textbooks, as the aforementioned time constraints can often prevent I would also like to thank my family whose love and support veterinary students from reading original literature. However, for sustains my existence, and Ira, we miss you so very much. You are those who may find the time, some original literature references always in our hearts and our memories. are also included in several chapters.
Finally, this book is a tribute to the veterinary students through- Accompanying resources for the text can be found on Elsevier’s out the world. It is these students who give pleasure, meaning, Evolve website. These include an online repository of the book’s and value to our teaching. Practice Questions and Clinical Correlations, as well as some relevant animations from Elsevier’s existing collection.
Students will appreci- Brad Klein ate the ability to access the Practice Questions as Practice Tests. vi Contents VetBooks.ir Section I: The Cell 8 Skeletal Muscle Receptor Organs, 86 Bradley G. Klein 1 The Molecular and Cellular Bases of Clinical Correlations: Physiological Regulation, 1 Femoral Nerve Mononeuropathy, 91 Steven R. Heidemann Obturator Nerve Paralysis in a Cow Post- Clinical Correlations: Calving, 91 Peripheral Edema, 27 Horse With Hyperkalemic Periodic Paralysis, 27 9 The Concept of Lower and Upper Motor Neurons and Their Malfunction, 93 2 Cancer: A Disease of Cellular Proliferation, Life Bradley G.
Klein Span, and Death, 29 Steven R.