Advances in Insect Physiology Volume 23 This Page Intentionally Left Blank Advances in Insect Physiology edited by P. D. EVANS Department of Zoology, The University Cambridge, England Volume 23 u ACADEMIC PRESS Harcourt Brace Jovanovich, Publishers London San Diego New York Sydney Toronto Tokyo Boston ACADEMIC PRESS LIMITED 24-28 Oval Road London NWl 7DX United States Edition published by ACADEMIC PRESS INC. San Diego, CA 92101 This book is printed on acid-free paper Copyright 0 1991 by ACADEMIC PRESS LIMITED All Rights Reserved No part of this book may be reproduced in any form by photostat, microfilm, or any other means, without written permission from the publishers A catalogue record for this book is available from the British Library ISBN 0-12424223-0 Typeset by Latimer Trend & Company Ltd, Plymouth and printed in Great Britain by St Edmundsbury Press Ltd, Bury St Edmunds, Suffolk Contributors S. A. Corbet Department of Zoology, Downing Street, Cambridge CB2 3EJ, UK C. P. Ellington Department of Zoology, Downing Street, Cambridge CB2 3EJ, UK R. E. Page, Jr Department of Entomology, University of California, Davis, C A 95616, USA M. P. Pener Department of Zoology, Hebrew University of Jerusalem, 91904 Jerusalem, Israel G. E. Robinson Department 05Entomology, University of Illinois, Urbanu, I L 61801, USA Retiring Editor V. B. Wigglesworth Advances in Insect Physiology, 1963-1990 Contents Contributors V Locust Phase Polymorphism and its Endocrine Relations M. P. PENER 1 A Fresh Look at the Arousal Syndrome of Insects S. A. CORBET 81 The Genetics of Division of Labour in Honey Bee Colonies R. E. PAGE, J R and G. E. ROBINSON 117 Aerodynamics and the Origin of Insect Flight C. P. ELLtNGTON 171 Subject Index 21 1 This Page Intentionally Left Blank Locust Phase Polymorphism and its Endocrine Relations M. P. Pener Department of Zoology, The Hebrew University of Jerusalem, 91904 Jerusalem, Israel 1 Insect polymorphism and its endocrine aspects 1 1.1 Polymorphism 1 1.2 Polymorphism and endocrine factors 3 2 Phase polymorphism 4 2.1 Locusts 4 2.2 Some other insects 7 3 Phase characteristics and related endocrine eEects 8 3.1 Morphology, morphometrics and anatomy 8 3.2 Colouration 12 3.3 Reproduction 21 3.4 Hopper development 26 3.5 Physiology, biochemistry and molecular biology 28 3.6 Cytology 30 3.7 Behaviour and activity 31 4 Endocrine organs, hormones and their role in phase transformation 37 4. I The corpora allata and juvenile hormone 37 4.2 The prothoracic (=ventral) glands and ecdysteroids 45 4.3 Neurosecretory cells, corpora cardiaca and neurohormones 48 5 Pheromones 49 6 Concluding remarks 52 References 55 Addenda 75 1 1.1 insect polymorphism and its endocrine aspects POLYMORPHISM The term polymorphism roughly means that distinct morphological differences, reflecting and often including physiological, behavioural and/or oecological differences, occur simultaneously or recurrently among conspecific individuals. Although the phenomenon is easily recognized and well known in many insect orders, its exact definition runs into difficulties. For example, most authors would not apply the term polymorphism to sexual ADVANCES IN INSECT PHYSIOLOGY VOL 23 ISBN 6 1 2 4 2 4 2 2 3 4 Copyrrghr 0 1991 Academic Press Limited AN rights of reproduction in any form reserved 2 M. P. PENER dimorphis