Insect Physiology and Biochemistry Second Edition 61771.indb 1 3/18/08 1:36:20 PM 61771.indb 2 3/18/08 1:36:20 PM Insect Physiology and Biochemistry Second Edition James L. Nation Department of Entomology and Nematology University of Florida Gainesville, Florida, U.S.A. 61771.indb 3 3/18/08 1:36:20 PM Cover photo by Rochelle Carlson Nation. CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2008 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed in the United States of America on acid-free paper 10 9 8 7 6 5 4 3 2 1 International Standard Book Number-13: 978-1-4200-6177-2 (Hardcover) This book contains information obtained from authentic and highly regarded sources Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the valid- ity of all materials or the consequences of their use. The Authors and Publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. 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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Nation, James L. Insect physiology and biochemistry / James L. Nation. p. ; cm. Includes bibliographical references and index. ISBN 978-1-4200-6177-2 (hardcover : alk. paper) 1. Insects--Physiology. I. Title. [DNLM: 1. Insects--physiology. QL 495 N277i 2008] QL495.N37 2008 571.1’57--dc22 2007045323 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com 61771.indb 4 3/18/08 1:36:21 PM Contents Preface..............................................................................................................................................vii Author ...............................................................................................................................................ix Chapter 1 Embryogenesis ...................................................................................................................................1 Chapter 2 Digestion ..........................................................................................................................................29 Chapter 3 Nutrition ...........................................................................................................................................69 Chapter 4 Integument .......................................................................................................................................91 Chapter 5 Hormones and Development ..........................................................................................................123 Chapter 6 Diapause .........................................................................................................................................161 Chapter 7 Intermediary Metabolism ..............................................................................................................177 Chapter 8 Neuroanatomy ................................................................................................................................205 Chapter 9 Neurophysiology ............................................................................................................................233 Chapter 10 Muscles ..........................................................................................................................................255 Chapter 11 Insect Flight ...................................................................................................................................279 Chapter 12 Sensory Systems ............................................................................................................................295 v 61771.indb 5 3/18/08 1:36:21 PM vi Insect Physiology and Biochemistry, Second Edition Chapter 13 Vision .............................................................................................................................................315 Chapter 14 Circulatory System.........................................................................................................................339 Chapter 15 Immunity .......................................................................................................................................367 Chapter 16 Respiration .....................................................................................................................................385 Chapter 17 Excretion ........................................................................................................................................417 Chapter 18 Pheromones ....................................................................................................................................447 Chapter 19 Reproduction ..................................................................................................................................483 Appendix .......................................................................................................................................511 Index ..............................................................................................................................................517 61771.indb 6 3/18/08 1:36:21 PM Preface In this second edition of Insect Physiology and Biochemistry, the chapter topics of the first edition have been retained and four new chapters have been added. Two of the new chapters (Chapter 6, Diapause, and Chapter 15, Immunity) cover topics and material not included in the first edition. Several anonymous reviewers of the first edition suggested that diapause and immunity needed to be included in any future edition, and I thank these reviewers for this advice. I also thank Dan Hahn for discussions of diapause and for valuable criticism and advice on Chapter 6, as well as Drion Boucias for helpful criticism and advice on immunity in Chapter 15. I take responsibility, however, for any shortcomings or errors in these chapters. Flight and Vision have been expanded into new chapters (Chapters 11 and 13, respectively) because of the large amount of new material discovered on the mechanics and dynamics of flight and color vision. New references have been added to each of the earlier chapters, as well as to the new chapters, so the book now includes nearly 1800 references to the literature. Published work in insect physiology and biochemistry seems to be expanding exponentially, and students and scien- tists need a guide to the literature. References from the 1960s, 1970s, and 1980s, and a few even earlier, have been retained because I think it is important to have an understanding of how insect physiology developed. I have been frustrated while reading books that supply few or no references to the literature with statements or views that I would like to check to see if my interpretation agrees with the author’s view. Just knowing facts usually is not as satisfying to me as knowing the evidence that supports why and how we know, or think we know, something. As in the first edition, my goal has been to provide a textbook that will serve advanced under- graduate and graduate students studying entomology and/or zoology, and for working scientists in molecular biology, genetics, endocrinology, virology, microbiology, and plant sciences who may have limited experience, but have an interest in the broader field of insect physiology. Teachers of college and university courses in insect physiology likely will have to be somewhat selective in how much of the book to cover in a one-semester course. I hope students will find the book interesting to read, and find it useful as a reference source as they go through graduate school and into one of the life sciences as a profession. I thank again those who aided and encouraged the first edition: Glenn Hall, Marie Nation Becker, Jon Harrison, Tom Miller, Kathy Milne, Christine Andreasen, and anonymous reviewers, because without their help I would not be doing a revision. I thank John Sulzycki for continuous encourage- ment with the first edition and this revision, and Pat Roberson, Jennifer Smith, and Zoe Tzanev at Taylor & Francis Group for editorial help with the revision. I appreciate the numerous illustrations that colleagues have let me use, and acknowledgment accompanies the appropriate illustrations in the book. Illustrations enhance a textbook, and I have tried to find and use good, illustrative ones. I thank Taylor & Francis Group for allowing several pages of color photos. I appreciate, as well, the permission that has been given to use many published figures. Finally, I thank my wife, Dorothy, for continued support and patience with this revision. James L. Nation Professor Emeritus, Department of Entomology and Nematology University of Florida, Gainesville vii 61771.indb 7 3/18/08 1:36:21 PM 61771.indb 8 3/18/08 1:36:21 PM Author James L. Nation, Ph.D., is professor of entomology at the University of Florida, Gainesville. He has a B.S. in entomology with a major in chemistry from Mississippi State University and a Ph.D. in entomology from Cornell University, where he specialized in insect physiology with a minor in insect biochemistry. He taught and conducted research at the University of Florida for 43 years, from 1960 until retirement in 2003. His primary teaching was in insect physiology, insect biochem- istry, and an honors course in global environmental issues. Research activities include nitrogen excretion, pheromones, cuticular hydrocarbons, and butterfly nutrition and rearing. He served as associate editor of Florida Entomologist from 1967 to 1969, as an editor of the Journal of Chemical Ecology from 1994 to 2000, and since 2004, as editor of Florida Entomologist, the official scientific journal of the Florida Entomological Society. He has continued occasional teaching at University of Florida, City College in Gainesville, and introduced a graduate level course in Insect Physiology at Florida A & M University in Tallahassee in 2006 (taught via interactive television). He is active in volunteer activities and in gardening. ix 61771.indb 9 3/18/08 1:36:22 PM
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