Insects at Low Temperature
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1 Insects at Low Temperature
2 Insects at Low Temperature Edited by Richard E. Lee, Jr. and David L. Denlinger Chapman and Hall New York and London
3 First published 1991 by Chapman and Hall 29 West 35th Street, New York, NY Published in Great Britain by Chapman and Hall, Ltd. 2-6 Boundary Row London SEI 8HN 1991 by Chapman and Hall Softcover reprint of the hardcover 1st edition 1991 All rights reserved. No part of this book may be reproduced in any way, or by any means, without permission in writing from the publisher. Library of Congress and British Library cataloging in publication data is available. ISBN DOl / ISBN (ebook)
4 Contents Preface ix Physiology of Insect Cold Hardiness 1. A Tribute to R. W. Salt Richard A. Ring and Paul W. Riegert 3 2. Principles of Insect Low Temperature Tolerance Richard E. Lee, Jr The Water Relations of Overwintering Insects Karl Erik Zachariassen Biochemistry of Cryoprotectants Kenneth B. Storey and Janet M. Storey Hemolymph Proteins Involved in Insect Subzero- Temperature Tolerance: Ice Nucleators and Antifreeze Proteins John G. Duman, Lei Xu, Lisa G. Neven, Donald Tursman, and Ding Wen Wu 94 Impact on Development and Survival 6. Cold Shock and Heat Shock David L. Denlinger, Karl H. Joplin, Cheng-Ping Chen, and Richard E. Lee, 1r Effects of Cold on Morphogenesis Frantisek Sehnal Relationship between Cold Hardiness and Diapause David L. Denlinger 174
5 vi / Contents 9. Thennoperiodism Stanley D. Beck 199 Species Adaptations 10. Winter Habitats and Ecological Adaptations for Winter Survival H. V. Danks Freezing Tolerance in the Goldenrod Gall Fly (Eurosta solidaginis) John G. Baust and Misako Nishino Behavioral and Physiological Adaptations to Cold in a Freeze-Tolerant Arctic Insect Olga Kukal Comparative Invertebrate Cold Hardiness Stephen H. Loomis Adaptations to Alpine and Polar Environments in Insects and Other Terrestrial Arthropods Lauritz Sj,1mme and William Block Overwintering of Freshwater Benthic Macroinvertebrates Mark W. Oswood, L. Keith Miller, and John G. Irons III 360 Practical Applications 16. Cryopreservation of Insect Gennplasm: Cells, Tissues, and Organisms Roger A. Leopold Cryobiology of Drosophila melanogaster Embryos Peter L. Steponkus, Stanley P. Myers, Daniel V. Lynch, Ronald E. Pitt, Ta-Te Lin, Ross J. MacIntyre, Stanley P. Leibo, and William F. RaIl Silkwonn Eggs at Low Temperatures: Implications for Sericulture Okitsugu Yamashita and Toshinobu Yaginuma Overwintering in Honey Bees: Implications for Apiculture Edward E. Southwick 446
6 Contents I vii 20. Implications of Cold Hardiness for Pest Management J. S. Bale Taxonomic Index Subject Index Contributors
7 Preface The study of insects at low temperature is a comparatively new field. Only recently has insect cryobiology begun to mature, as research moves from a descriptive approach to a search for underlying mechanisms at diverse levels of organization ranging from the gene and cell to ecological and evolutionary relationships. Knowledge of insect responses to low temperature is crucial for understanding the biology of insects living in seasonally varying habitats as well as in polar regions. It is not possible to precisely define low temperature. In the tropics exposure to C may induce chill coma or death, whereas some insects in temperate and polar regions remain active and indeed even able to fly at O C or below. In contrast, for persons interested in cryopreservation, low temperature may mean storage in liquid nitrogen at C. In the last decade, interest in adaptations of invertebrates to low temperature has risen steadily. In part, this book had its origins in a symposium on this subject that was held at the annual meeting of the Entomological Society of America in Louisville, Kentucky, USA in December, However, the emergence and growth of this area has also been strongly influenced by an informal group of investigators who met in a series of symposia held in Oslo, Norway in 1982, in Victoria, British Columbia, Canada in 1985 and in Cambridge, England in Another is scheduled for Binghamton, New York, USA (1990). The literature on insect cold tolerance is widely scattered, thus making it difficult for students and new workers to gain quick access to this field. Currently no single volume exists on the topic of insects at low temperature. Furthermore, in recent years fundamental tenets of insect cold-hardiness have been questioned and new hypotheses proposed. This book is designed to describe the current state of knowledge in this field as well as suggest areas for future study. The first chapter pays tribute to R. W. Salt for his pioneering work on insect cold-hardiness that began in the 1930s and continued into the 1970s. It is humbling to reflect that the fundamental questions and problems raised by Salt--chilling and freezing injury, the nature of supercooling and nucleation, action of cryopro-
8 x / Preface tectants, environmental regulation of cold-hardening, and the significance of cold tolerance for the conrol of insect pests-remain the primary focus of our research today. An introductory section (Chapters 2-5) defines basic concepts and summarizes our understanding of biochemical and physiological principles of insect coldhardiness. Section II (Chapters 6-9) integrates the impact of low temperature on development and survival, and includes discussions of cold shock and heat shock, morphogenesis, diapause, and thermoperiodism. Chapters introduce the diversity of ecological and physiological adaptations for both survival and activity at low temperature in intensely studied species and for groups of insects and related arthropods living in polar and alpine regions. The final section (Chapters 16-20) examines practical applications of the study of insect cold-hardiness and cryopreservation. Chapters are devoted to recent advances in the cryopreservation of germplasm, cells,and entire insects, particularly Drosophila, and to the implications of low temperature in the management of beneficial insects (honey bees and silkmoths) and pest species. Finally we thank our colleagues for their contributions and Gregory W. Payne, Science Editor for Chapman and Hall, Inc. for his editorial assistance. We gratefully acknowledge the National Science Foundation, the National Institutes of Health, and the United States Department of Agriculture for their support of research in our laboratories. R.E. Lee Hamilton, Ohio D.L. Denlinger Columbus, Ohio October, 1989
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