Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information The Insects Structure and Function FIFTH EDITION The Insects has been the standard textbook in the field since the first edition was published over 40 years ago. Building on the strengths of Chapman’s original text, this long-awaited new edition has been revised and expanded by a team of eminent insect physiologists, bringing it fully up to date for the molecular era. The chapters retain the successful structure of the earlier editions, focusing on particular functional systems rather than on taxonomic groups and making it easy for students to delve into topics without extensive knowledge of taxonomy. The focus is on form and function, bringing together basic anatomy and physiology and examining how these relate to behavior. This, combined with nearly 600 clear illustrations, provides a comprehensive understanding of how insects work. Now also featuring a richly illustrated prologue by George McGavin, this is an essential text for students, researchers and applied entomologists alike. R. F. Chapman (1930–2003) was an eminent insect physiologist and Professor in the Division of Neurobiology at the University of Arizona. His first four editions of The Insects have formed the standard text in the field for more than 40 years. Stephen J. Simpson is ARC Laureate Fellow in the School of Biological Sciences and Academic Director of the Perkins Centre for the study of obesity, diabetes and cardiovascular disease at the University of Sydney. His core research aims are to understand swarming in locusts and to develop and implement an integrative framework for studying nutrition. In 2012 he was awarded the Wigglesworth Medal from the Royal Entomological Society of London. Angela E. Douglas is Daljit S. and Elaine Sarkaria Professor of Insect Physiology and Toxicology at Cornell University, New York. Her research and teaching is motivated by the mechanisms underlying insect function, and her core research interests are the overlapping topics of insect nutrition and interactions between insects and beneficial microorganisms. She is a Fellow of The Royal Entomological Society and The Entomological Society of America. © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information The Insects Structure and Function FIFTH EDITION R. F. CHAPMAN Formerly of the University of Arizona, USA Edited by STEPHEN J. SIMPSON The University of Sydney, Australia ANGELA E. DOUGLAS Cornell University, New York, USA © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information CAMBRIDGE UNIVERSITY PRESS Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sa˜o Paulo, Delhi, Mexico City Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521113892 © Cambridge University Press 1998, 2013 This publication is in copyright. Subject to statutory exception and to the provisions of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published by Edward Arnold 1969 Second edition 1971, 6th printing 1980 Third edition 1982, 5th printing 1991 Fourth edition published by Cambridge University Press 1998, 7th printing 2011 Fifth edition 2013 Printed in the United States of America A catalogue record for this publication is available from the British Library Library of Congress Cataloguing in Publication data Chapman, R. F. (Reginald Frederick) The insects : structure and function / R. F. Chapman. – 5th edition / edited by Stephen J. Simpson, Angela E. Douglas. pages cm Includes bibliographical references and indexes. ISBN 978-0-521-11389-2 1. Insects. I. Simpson, Stephen J. II. Douglas, A. E. (Angela Elizabeth), 1956– III. Title. QL463.C48 2013 595.7–dc23 2012018826 ISBN 978-0-521-11389-2 Hardback Cambridge University Press has no responsibility for the persistence or accuracy of URLs for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information CONTENTS List of contributors ix 4 Nutrition 81 Preface xi Introduction 81 Acknowledgments xii 4.1 Required nutrients 82 Prologue xiii 4.2 Balance of nutrients 87 4.3 Nutritional effects on growth, development, Part I The head, ingestion, utilization reproduction and lifespan 95 and distribution of food 4.4 Contribution of symbiotic microorganisms to insect nutrition 99 1 Head 3 Summary 104 Introduction 3 1.1 Head 4 5 Circulatory system, blood and the 1.2 Neck 9 immune system 107 1.3 Antennae 10 Introduction 107 Summary 13 5.1 The circulatory system 108 5.2 Circulation 113 2 Mouthparts and feeding 15 5.3 Hemolymph 117 Introduction 15 5.4 Hemocytes 124 2.1 Ectognathous mouthparts 16 Summary 129 2.2 Mechanics and control of feeding 22 2.3 Regulation of feeding 34 6 Fat body 132 2.4 Other consequences of Introduction 132 feeding 37 6.1 Fat body structure and development 133 2.5 Head glands 37 6.2 Storage and utilization of energy Summary 43 and nutrients 137 6.3 Function as an endocrine organ and 3 Alimentary canal, digestion and nutritional sensor 142 absorption 46 Summary 144 Introduction 46 3.1 The alimentary canal 47 Part II The thorax and locomotion 3.2 Digestion 59 3.3 Absorption 72 7 Thorax 149 3.4 The alimentary tract as an immunological Introduction 149 organ 77 7.1 Segmentation of the thorax 150 Summary 78 7.2 Morphology of the thorax 151 © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information vi Contents 7.3 Muscles of the thorax 155 12 Reproductive system: male 282 Summary 155 Introduction 282 12.1 Anatomy of the internal reproductive 8 Legs and locomotion 157 organs 283 Introduction 157 12.2 Spermatozoa 286 8.1 Structure of the legs 158 12.3 Transfer of sperm to 8.2 Walking and running 166 the female 292 8.3 Other mechanisms of terrestrial 12.4 Other effects of mating 306 locomotion 173 Summary 310 8.4 Aquatic locomotion 180 8.5 Other uses of legs 185 13 Reproductive system: female 313 Summary 189 Introduction 313 13.1 Anatomy of the internal reproductive 9 Wings and flight 193 organs 314 Introduction 193 13.2 Oogenesis 317 9.1 Structure of the wings 194 13.3 Ovulation 332 9.2 Form of the wings 204 13.4 Fertilization of the egg 333 9.3 Movement of the wings 207 13.5 Oviposition 334 9.4 Wing kinematics 214 Summary 343 9.5 Aerodynamic mechanisms 221 9.6 Power for flight 223 14 The egg and embryology 347 9.7 Sensory systems for flight control 225 Introduction 347 Summary 230 14.1 The egg 348 14.2 Embryogenesis 357 10 Muscles 233 14.3 Alternative strategies of acquiring nutrients Introduction 233 by embryos 380 10.1 Structure 234 14.4 Sex determination 388 10.2 Muscle contraction 242 14.5 Parthenogenesis 390 10.3 Regulation of muscle contraction 244 14.6 Pedogenesis 392 10.4 Energetics of muscle contraction 252 Summary 393 10.5 Muscular control in the intact insect 254 10.6 Changes during development 257 15 Postembryonic development 398 Summary 263 Introduction 398 15.1 Hatching 399 Part III The abdomen, reproduction 15.2 Larval development 403 and development 15.3 Metamorphosis 417 15.4 Control of postembryonic 11 Abdomen 269 development 436 Introduction 269 15.5 Polyphenism 443 11.1 Segmentation 270 15.6 Diapause 448 11.2 Abdominal appendages and outgrowths 273 Summary 454 Summary 280 © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-0-521-11389-2 - The Insects: Structure and Function: Fifth Edition R. F. Chapman Edited by Stephen J. Simpson and Angela E. Douglas Frontmatter More information Contents vii Part IV The integument, gas exchange 19 Thermal relations 588 and homeostasis Introduction 588 19.1 Body temperature 589 16 Integument 463 19.2 Thermoregulation 595 Introduction 463 19.3 Performance curves 598 16.1 Epidermis 464 19.4 Behavior and survival at low 16.2 The cuticle 469 temperatures 600 16.3 Chemical composition of the cuticle 473 19.5 Activity and survival at high 16.4 Types of cuticles 483 temperatures 607 16.5 Molting 488 19.6 Acclimation 609 16.6 Cuticle formation 493 19.7 Cryptobiosis 611 16.7 Functions of the integument 497 19.8 Temperature and humidity receptors 611 Summary 498 19.9 Temperature-related