Table Of ContentInsect
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
