Insect flight PDF
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Insect Flight Editors Graham J. Goldsworthy Professor and Head Department of Biology Birkbeck College University of London London, England Colin H. Wheeler Research Officer Department of Biology Birkbeck College University of London London, England Boca Raton London New York CRC Press, Inc. CRC Press is an imprint of the TBayolocra & RFraantcoisn G,r oFuplo, arni dinaforma business First published 1989 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1989 by CRC Press, Inc. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works 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 validity 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. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http://www.copyright. com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not- for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. 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 Insect flight/editors, Graham J. Goldsworthy, Colin H. Wheeler. p. cm. Bibliography: p. Includes index. ISBN 0-8493-4969-9 1. Insects--Flight. I. Goldsworthy, Graham J. II. Wheeler, Colin H. QL496.7.157 1989 595.7’01852--dc19 88-832 A Library of Congress record exists under LC control number: 88000832 Publisher’s Note The publisher has gone to great lengths to ensure the quality of this reprint but points out that some imperfections in the original copies may be apparent. Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-1-315-89448-5 (hbk) ISBN 13: 978-1-351-07358-5 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com PREFACE Insects are the most numerous class of animals on earth, both in terms of their numbers and their variety. There are more than 800,000 recognized species, with between 1 and 10 million estimated species yet to be classified! Their tremendous success is attested by their exploitation of environmental niches ranging from deserts to ice fields, from freshwater streams to hot springs. What is it about the design of insects that has made them so successful? Is it their rigid, impermeable exoskeleton, their ability to fly, their high reproductive potential, or their small size? Perhaps it is the adaptability of their behavior and physiology to changing environmental conditions. It is likely to be a combination of all these things, but in these chapters we will focus our attention on only one: their ability to fly. The adoption of a rigid exoskeleton certainly meant that many insects can crawl, jump, run, and even swim with great efficiency. Such forms of locomotion are undoubtedly of great benefit in escaping from predators, capturing prey, or searching for food or mates. However, the development of flight placed many insects beyond the reach of competitors and undoubtedly played a great part in the success of insects as terrestrial animals. This ability to fly was established by insects over capturing prey, or searching for food or mates. However, the development of flight placed many insects beyond the reach of competitors and undoubtedly played a great part in the success of insects as terrestrial animals. This ability to fly was established by insects over 300 million years ago as part of a general pattern of winged organization which does not appear to have changed greatly since that time, suggesting that it proved to be a phenomenally successful design almost from the outset. Flight activity may be relatively trivial or routine, associated with feeding or escape, or it may form part of a complex and often spectacular pattern of behavior associated with courtship, dispersal, or migration. No student of biology should fail to wonder at the sheer grace and beauty of insects in flight, whether as individual insects or as members of a swarm. But the study of insects in flight is no idle curiosity of ivory-towered scientists. For, while we believe that scientific curiosity is worthwhile in itself, there is nevertheless immense practical value in the study of insect flight, and man's natural spirit of inquiry should therefore be encouraged and not suppressed in times of economic restraint. There is no doubt that insect material can be used to good effect as "models" to gain insight into basic features of animal physiology, behavior, and ecology, but perhaps even more importantly, we need to know more about insects, at all levels of their existence. We have an enigmatic relationship with the insects; on the one hand they are essential for the pollination of many of our crops, but on the other they are our most successful competitors on this planet. Not only do they eat our crops, fabrics, and timbers, but they spread disease among ourselves and our animals. Again, their success in all these activities is enhanced by the ability of many to fly. How can we fight back? During flight, the general behavior of the insect, its movement in space, and the coor- dinated movements of the wings, are dependent on energy production (metabolism), res- piration, transport of metabolites, muscular contraction, hormonal and sensory control, and mechanical or aerodynamic principles. The chapters of this book have been specially written by internationally recognized authorities in these facets of insect flight. The discussions contained therein extend from studies at the molecular level to those of populations of whole animals. In the two concluding chapters, the implications of flight for insect pest control are outlined, and the prospects for better control strategies are discussed. It is ironic that studies of insect flight, for many scientists born of a fascination with the insect itself, should be used to further methods of pest control, but at a time when insecticide resistance is still a major problem, better and more efficient control strategies are urgently required. However, such strategies must be more selective if we are to avoid the wide-scale ecological disasters which can be caused by the profligate use of nonspecific control agents. Many insect species are beneficial to man, and we must also be wary of the short- and long- term effects of control agents on other organisms and on ourselves. Insecticides directed at particular aspects of an insect's physiology, behavior, or ecology, formulated and used with a full understanding of that biology, may provide the necessary means for controlling specific pests in a highly selective manner. Such developments will depend on in-depth knowledge of all aspects of insect biology, including flight. The challenges are here for all scientists whether they be interested in mankind, insects, or both. THE EDITORS Graham Goldsworthy, Ph.D., F.R.E.S., is Professor and Head of Biology at Birkbeck College in the University of London, U.K. Professor Goldsworthy obtained both his first degree (1966) and his doctorate (1969) from the University of Sheffield. These were followed by 2 years of research in the University of Sussex, before taking up a lectureship in the University of Hull. He was appointed as Reader in Comparative Endocrinology in the University of Hull in 1978, and Professor of Biology in the University of London in 1986. Professor Goldsworthy has been on the Editorial Board of the Journal of Insect Physiology (1978 to 1984) and is currently the senior editor of Physiological Entomology for the Royal Entomological Society of London, was the Publications Officer of the Society for Experi- mental Biology (1979 to 1983), and has been a member of Council of the European Society for Comparative Endocrinology (1981 to 1986). Colin H. Wheeler, Ph.D., is a Research Officer at Birkbeck College in the University of London, U.K. Dr. Wheeler obtained his first degree (1977) and his doctorate (1981) from the University of Hull. After gaining his doctorate, Dr. Wheeler spent a year working with Professor Beenakkers in the University of Utrecht, during which time he became interested in lipoprotein Upases in the flight muscle of locusts. He moved to Birkbeck in 1986. CONTRIBUTORS D. J. Aidley, Ph.D. David P. Giles, Ph.D. School of Biological Sciences Head of Department of Entomology and University of East Anglia Nematology Norwich, England Agrochemical Research Schering AG Thomas C. Baker, Ph.D. Berlin, West Germany Professor Department of Entomology A. R. Jutsum, Ph.D., F.R.E.S. University of California Head of Entomology Riverside, California ICI Agrochemicals Jealott's Hill Research Station D. J. Candy, Ph.D. Bracknell Department of Biochemistry Berkshire, England University of Birmingham Birmingham, England Wolfram Kutsch, Prof. Dr. Professor Timothy M. Casey, Ph.D. Faculty of Biology Professor of Zoology University of Konstanz Department of Entomology and Economic Konstanz, West Germany Zoology Bernhard Mohl, Priv. Doz. Dr. Cook College Wissenschaftlicher Mitarbeiter Rutgers University Zoology Institute New Brunswick, New Jersey University of Saarland Saarbriicken, West Germany Richard John Cooter, Ph.D. Head of Physiology and Behavior Section W. Nachtigall, Prof. Dr. Applied Ecology Department Professor Overseas Development and Natural Department of Zoology Resources Institute University of Saarland Chatham, England Saarbrucken, West Germany Hugh Dingle, Ph.D. Mary Ann Rankin Professor Department of Zoology Department of Entomology University of Texas University of California Austin, Texas Davis, California Thomas D. Seeley, Ph.D. Fred C. Dyer, Ph.D. Associate Professor Assistant Professor Section of Neurobiology and Behavior Department of Zoology Cornell University Michigan State University Ithaca, New York East Lansing, Michigan Colin H. Wheeler, Ph.D. A. G. Gatehouse, Ph.D. Research Officer Senior Lecturer Department of Biology School of Biological Sciences Birkbeck College University College of North Wales University of London Bangor, Gwynedd, Wales London, England To John Guest Phillips, FRS (1933 — 1987) A good friend and mentor TABLE OF CONTENTS Chapter 1 Mechanics and Aerodynamics of Flight 1 W. Nachtigall Chapter 2 Structure and Function in Flight Muscle 31 D. J. Aidley Chapter 3 Development of the Flight Motor Pattern 51 Wolfram Kutsch Chapter 4 Sense Organs and the Control of Flight 75 Bernhard Mohl Chapter 5 The Evolution and Significance of Migratory Flight 99 Hugh Dingle Chapter 6 Genes, Environment, and Insect Flight 115 A. G. Gatehouse Chapter 7 Hormonal Control of Flight 139 Mary Ann Rankin Chapter 8 Swarm Flight Behavior in Flies and Locusts 165 Richard John Cooler Chapter 9 Orientation Foraging in Honey Bees 205 Fred C. Dyer and Thomas D. Seeley Chapter 10 Pheromones and Flight Behavior 231 T. C. Baker Chapter 11 Oxygen Consumption During Flight 257 Timothy M. Casey Chapter 12 Mobilization and Transport of Fuels to the Flight Muscles 273 Colin H. Wheeler Chapter 13 Utilization of Fuels by the Flight Muscles 305 D. J. Candy Chapter 14 Problems in the Control of Flying Insect Pests 321 D. P. Giles and A. R. Jutsum Chapter 15 Prospects for Better Control Strategies 337 A. R. Jutsum and D. P. Giles Index 359
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