Adtmiicrs in INSECT PHYSIOLOGY VOLUME 6 This Page Intentionally Left Blank Advances in Insect Physiology Edited by J. W. L. BEAMENT, J. E. TREHERNE and V. B. WICGLESWORTH Deparment of Zoology, The Unfversify,Cambridge, Emgland VOLUME 6 I ’ I I AN ACADEMIC PRESS REPLICA REPRINT 1969 This is an Academic Press Replica Reprint reproduced directly from the pages of a title for which type, plates, or film no longer exist. Although not up to the standards of the original, this method of reproduction makes it possible to provide copies of books which otherwise would be out of print. ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Roud. London N W I U.S. Edition published by ACADEMIC PRESS. INC I I I FiAh Avenue. New York. New York lo003 Copyright 9 I!%Y By Academic Press Inc. (London) Ltd. AN Rights Reserved NO PART OF THIS BOOK MAY BE REPRODUCED I N ANY FORM BY PHOTOSTAT, MICROFILM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM THE PUBLISHERS Library of Congress Cutulog Card Numbcr: 63-14039 SBN: 12-024206-0 Printed in Great Britain by The Whitefriars Press Ltd., London and Tonbridge 8 0 8 1 82 9 8 7 6 5 4 3 List of Contributors to Volume 6 E. ASAHINA, The Institute of Low Temperahwe Science, Hokkuido University, Sapporo, Japun (p. 1) A. D . CARLSON, Department of Biological Sciences, State University of New York a t Stony Brook, Stony Brook. New York, L?S.As.(p. 5 1) JOHN S . EDWARDS,Deprtment of Zoology, Unfversity of Washington, Seattle, Washington, LI.S.A. (p. 97) RUDOLF HARMSEN, Biology Department, Queen% Univeraity. Kingston, Canuda (p. 139) P.N. R . USHERWOOD,Department of Zoology, University of Glasgow, Glosgow, Scothnd (p. 203) IRMGARD ZIECLER, Botanical Institute, Darmstadt institute of Technology, Darmstadt, Germany (p. 139) V Acknowledgements The Editors wish to thank the publishers of Arch. ital. Biol., Wilhelm Roux Arch., J. Insect Physiol., Comp. Biochem. Physiol., Nature, Lond., Biol. Bull., J. exp. Biol. and J. Cell Biol. for their $ind permission to reproduce figures from their publications. The authors in this volume have been instructed to obtain necessary permissions for use of published figures and the Editors sincerely regret the omission in case any of the sources were overlooked. J. W. L. Beament J. E. Treherne V.B. Wigglesworth Contents LIST OF CONTRIBUTORS TO VOLUME 6 . . . . . . . . . . v FROST RESISTANCE IN INSECTS . E ASAHINA I . Introduction . . . . . . . . . . . . . . . . . . . I1. Initiation of Freezing . . . . . . . . . . . . . . . . A Spontaneous Freezing . . . . . . . . . . . . . . . B . Inoculation by External Ice Crystals . . . . . . . . . . 111. The Freezing Process . . . . . . . . . . . . . . . . A. Freezing of the Blood . . . . . . . . . . . . . . . B . Types of Cell Freezing . . . . . . . . . . . . . . . C . Possible Process of Freezing in an Intact Insect . . . . . . . D Amount of Ice Formed in an Insect . . . . . . . . . . IV. Frost !Jury . . . . . . . . . . . . . . . . . . . A . Process of Injury . . . . . . . . . . . . . . . . . B Nature of Injury . . . . . . . . . . . . . . . . . C Injury Resulting from Extended Freezing Periods . . . . . . D. Effect of Freezing on the Termination of Diapause . . . . . V . Mechanisms of Frost Resistance . . . . . . . . . . . . . A Avoidagce of Intracellular Freezing . . . . . . . . . . . B Resistance t o Extracellular Freezing . . . . . . . . . . C Process of Frost Hardening . . . . . . . . . . . . . Vi. Freezing Resistance in Insects at Artificial very Low Temperatures . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 2 3 4 4 7 8 1 2 1 4 1 4