Table Of ContentContributors
MOISES AGOSIN
FRANCISCO J. AYALA
MURRAY S. BLUM
P. S. CHEN
G. MICHAEL CHIPPENDALE
W. C. DAUTERMAN
ROGER G. H. DOWNER
ERNEST HODGSON
A. E. NEEDHAM
R. D. O'BRIEN
A. GLENN RICHARDS
LYNN M. RIDDIFORD
WENDELL L. ROELOFS
JAMES W. TRUMAN
NEVIN WEAVER
Biochemistry
of Insects
EDITED BY
Morris Rockstein
Department of Physiology and Biophysics
School of Medicine
University of Miami
Coral Gables, Florida
ACADEMIC PRESS New York San Francisco London 1978
A Subsidiary of Harcourt Brace )ovanovich, Publishers
To SUZI
and
MADY
COPYRIGHT © 1978, BY ACADEMIC PRESS, INC.
ALL RIGHTS RESERVED.
NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR
TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC
OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY
INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT
PERMISSION IN WRITING FROM THE PUBLISHER.
ACADEMIC PRESS, INC.
Ill Fifth Avenue, New York, New York 10003
United Kingdom Edition published by
ACADEMIC PRESS, INC. (LONDON) LTD.
24/28 Oval Road, London NW1 7DX
Library of Congress Cataloging in Publication Data
Main entry under title:
Biochemistry of insects.
Includes bibliographies and index.
1. Insects—Physiology. I. Rockstein, Morris.
[DNLM: 1. Insects—Metabolism. 2. Insects—Physiol
ogy. QX500.3B615]
QL495.B56 595.7ΌΓ92 77-11221
ISBN 0-12-591640-X
PRINTED IN THE UNITED STATES OF AMERICA
To SUZI
and
MADY
COPYRIGHT © 1978, BY ACADEMIC PRESS, INC.
ALL RIGHTS RESERVED.
NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR
TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC
OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY
INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT
PERMISSION IN WRITING FROM THE PUBLISHER.
ACADEMIC PRESS, INC.
Ill Fifth Avenue, New York, New York 10003
United Kingdom Edition published by
ACADEMIC PRESS, INC. (LONDON) LTD.
24/28 Oval Road, London NW1 7DX
Library of Congress Cataloging in Publication Data
Main entry under title:
Biochemistry of insects.
Includes bibliographies and index.
1. Insects—Physiology. I. Rockstein, Morris.
[DNLM: 1. Insects—Metabolism. 2. Insects—Physiol
ogy. QX500.3B615]
QL495.B56 595.7ΌΓ92 77-11221
ISBN 0-12-591640-X
PRINTED IN THE UNITED STATES OF AMERICA
List of Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
MOISES AGO SIN (93), Department of Zoology, University of Georgia,
Athens, Georgia
FRANCISCO J. AYALA (579), Department of Genetics, Agricultural Exper
iment Station, College of Agricultural and Environmental Sci
ences, University of California, Davis, California
MURRAY S. BLUM (465), Department of Entomology, University of Geor
gia, Athens, Georgia
P. S. CHEN (145), Institute of Zoology and Comparative Anatomy, Univer
sity of Zurich, Zurich, Switzerland
G. MICHAEL CHIPPENDALE (1), Department of Entomology, College of
Agriculture, University of Missouri-Columbia, Columbia, Mis
souri
W. C. DAUTERMAN (541), Department of Entomology, North Carolina
State University, Raleigh, North Carolina
ROGER G. H. DOWNER (57), Department of Biology, Faculty of Science,
University of Waterloo, Waterloo, Ontario, Canada
ERNEST HODGSON (541), Department of Entomology, North Carolina
State University, Raleigh, North Carolina
A. E. NEEDHAM (233), Department of Zoology, Oxford University, Ox
ford, England
R. D. O'BRIEN (515), Section of Neurobiology and Behavior, Langmuir
Laboratory, Cornell University, Ithaca, New York
A. GLENN RICHARDS (205), Department of Entomology, Fisheries, and
Wildlife, University of Minnesota Twin Cities, St. Paul, Minnesota
XI
xii List of Contributors
LYNN M. RIDDIFORD (307), Department of Zoology, University of Wash
ington, Seattle, Washington
WENDELL L. ROELOFS (419), Department of Entomology,
Entomology-Plant Pathology Laboratory, New York State Ag
ricultural Experiment Station, Cornell University, Geneva, New
York
JAMES W. TRUMAN (307), Department of Zoology, University of Wash
ington, Seattle, Washington
NEVIN WEAVER (359, 391), Department of Biology, University of
Massachusetts-Boston, Harbor Campus, Boston, Massachusetts
Preface
The science of biochemistry may be viewed as microanatomy studied at
its ultimate level. Indeed, knowledge of an organism's intimate molecular
makeup, and of its component parts as well, adds an important dimension to
our understanding of its life processes.
Insect biochemistry as we know it today evolved from earlier efforts of
such biologists as Keilin, Wigglesworth, and Fraenkel. Later studies by
Chadwick, Dethier, Williams, and Richards (and by their students)
broadened the range and extended the depth of our knowledge of insectan
biochemistry, especially over the past two decades. More recently, increas
ing numbers of biochemists have exploited the wide versatility of this largest
single taxonomic group for basic biochemical studies per se. Taken as a
whole, their combined published researches in insectan biochemistry have
dictated the need for a summary volume such as this edited work, which was
intended as an in-depth updating of the only prior substantive book, by
Darcy Gilmour, published in 1961. The extensive coverage provided by this
book reflects the progress made since that time in insect biochemistry as a
generic science.
Designed to serve as a basic textbook in field, this volume should be equally
useful as an auxiliary text for most relevant courses in insect biology,
particularly insect physiology, insect ecology, insect control, and economic
entomology. To this end, the authors have attempted to present the material
at the level of a teaching aid. But in a few chapters of highly technical subject
matter, has this not been completely feasible. In any case, a background
in general biochemistry is a prerequisite. The book should also serve as an
important reference source for the advanced student, the research scientist,
and the professional entomologist seeking authoritative details of relevant
areas of subject matter. Most chapters include two lists of references. One is
a list of general references intended particularly for students, and includes
books, chapters in books, and review articles. The second is a detailed list of
specific, original publications intended for advanced students and research
scientists. In some cases, where the literature is more limited, only one list
of references, mostly general in nature, has been included.
xiii
xiv Preface
As in any general work in which the names of insects are cited, the problem
of synonymy arises. The knowledgeable entomologist will experience some
dismay at the occasional use of different names for the same insect. These
differences naturally are due to the fact that, when the need arises, taxonomists
must correct previously employed scientific names. The average reader will
not view this as a problem. To the purist, the insect taxonomist in particular,
such encounters will prove understandably painful.
The Editor acknowledges with thanks the advice of his former mentor and
present colleague, Dr. Glenn Richards, for his advice in the early stages of
planning the scope and contents of this book. To all of the contributors goes
my sincere appreciation for their cooperation in helping to maintain the
originally planned schedule for the publication of this compendious work.
Morris Rockstein
1
The Functions of Carbohydrates
in Insect Life Processes
G. MICHAEL CHIPPENDALE
I. Introduction 2
II. Classification of Insect Carbohydrates 3
III. Carbohydrates in Insect Nutrition 4
A. Larval Carbohydrate Requirements 5
B. Adult Carbohydrate Requirements 7
C. Carbohydrate Growth Factors 8
IV. Digestion of Carbohydrates 11
A. Common Digestive Carbohydrases 11
B. Special Digestive Carbohydrases 13
V. Absorption of Carbohydrates 15
VI. Excretion and Detoxication 16
A. Regulation of Carbohydrate Excretion 16
B. Glycosides in Detoxication 18
VII. Carbohydrates in Structural Components and
Pigments 19
A. Chitin 19
B. Glycosides and Sclerotization 20
C. Glycosidic Pigments and Cardiac Glycosides 20
D. Glycosaminoglycans in Connective Tissues and
Cement 22
VIII. Carbohydrates in Insect Metabolism 23
A. Synthesis, Nature, and Storage of Carbohydrate
Reserves 23
B. Mobilization of Carbohydrate Reserves 29
C. Glycolysis and the Pentose Phosphate Pathway 31
D. Regulation of Carbohydrate Metabolism 32
IX. Carbohydrates in Metamorphosis 34
X. Carbohydrates in Flight Muscles 37
XI. Carbohydrates in Reproduction and Embryonic
Development 42
1
2 G. Michael Chippendale
A. Male and Female Reproductive Systems 42
B. Embryonic Development 44
XII. Carbohydrates in Diapause and Coldhardiness 45
A. Diapause Maintenance 45
B. Polyols in Coldhardiness 46
XIII. Carbohydrates and Insect Behavior 49
A. Carbohydrates in Feeding Behavior 49
B. Glycosides in Exocrine Glands 52
XIV. Conclusion 52
General References 53
References for Advanced Students and Research Scientists . . 54
I. INTRODUCTION*
Carbohydrates, along with proteins and lipids, form the principal classes
of organic compounds that are found in insects and other organisms. Carbo
hydrates contribute to the structure and functions of all insect tissues, and
can be found in the nuclei, cytoplasm, and membranes of cells, as well as in
the extracellular hemolymph and supporting tissues. The term 4'carbohy
drate" is applied to polyhydroxy aldehydes and ketones and their deriva
tives. The classification of carbohydrates as monosaccharides, disac-
charides, oligosaccharides, and polysaccharides according to the number of
sugar residues found in the molecule has been well established. Insect
carbohydrates are present both in free form and combined with other mole
cules, including purines, pyrimidines, proteins, and lipids. Carbohydrates
are involved at all levels of cellular organization.
In general, carbohydrate metabolism in insects is similar to that found in
vertebrates. However, insects, unlike vertebrates, possess an exoskeleton
that is rich in the aminopolysaccharide, chitin. They also contain a disac-
charide, trehalose, which acts as a storage form of glucose. Special features
of insect carbohydrate metabolism, therefore, center around the synthesis
and hydrolysis of these compounds.
The function of carbohydrates in regulating and maintaining the life pro
cesses of insects is examined in this chapter. Included are accounts of
dietary carbohydrate requirements, the digestion and absorption of carbo
hydrates, the involvement of carbohydrates in excretory processes, the
nature of structural carbohydrates and carbohydrate pigments, and the
metabolic interconversions and breakdown of carbohydrates. In addition,
the origin, nature, and fate of carbohydrates in specific tissues during
*The following standard abbreviations are used in the text: ADP, adenosine 5'-diphosphate;
ATP, adenosine 5'-triphosphate; CoA, coenzyme A; cyclic AMP, adenosine 3':5'-
monophosphate; DNA, deoxyribonucleic acid; FAD, flavin-adenine dinucleotide; FMN, flavin
mononucleotide; NAD+, NADP+, nicotinamide-adenine dinucleotide (phosphate); RNA,
ribonucleic acid; UDP, uridine 5'-diphosphate; UTP, uridine 5'-triphosphate.
