Insect Physiology the Principles of Insect Physiology

THE PRINCIPLES OF INSECT PHYSIOLOGY THE PRINCIPLES OF INSECT PHYSIOLOGY

by V. B. WIGGLESWORTH C.B.E., M.D., F.R.S.

EMERITUS PROFESSOR OF BIOLOGY IN THE UNIVERSITY OF CAMBRIDGE; FORMERL Y DIRECTOR, AGRICULTURAL RESEARCH COUNCIL UNIT OF INSECT PHYSIOLOGY

Seventh Edition

With 412 illustrations

LONDON NEW YORK CHAPMAN AND HALL First published in 1939 by Methuen and Co. Ltd Second edition 1944 Third edition 1947 Fourth edition, revised, 1951 Fifth edition, with addenda, 1953 Sixth edition, revised, 1965 Reprinted once Seventh edition 1972 published by Chapman and Hall Ltd 11 New Fetter Lane London EC4P 4EE First published as a Science Paperback 1ffl2 Published in the USA by Chapman and Hall 733 Third Avenue New York NY 10017

ISBN 978-0-41 2-24660-9 ISBN 978-94-009-5973-6 (eBook) DOl 10. I 007 I 978-94-009-5973-6 This paperback edition is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, re-sold, hired out, or otherwise circulated without the publisher's prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser.

All rights reserved. No part of this book may be reprinted, or reproduced or utilized in any form or by any electronic, mechanical or other means, now known or hereafter in vented, including photocopying and recording, or in any information storage and retrieval system, without per mission in writing from the Publisher. PREFACE INSECTS PROVIDE an ideal medium in which to study all the problems of physiology. But if this medium is to be used to the best advantage, the principles and peculiarities of the insect's organization must be first appreciated. It is the purpose of this book to set forth these principles so far as they are understood at the present day. There exist already many excellent text-books of general ento• mology; notably those of Imms, Weber, and Snodgrass, to mention only the more recent. But these authors have necessarily been preoccupied chiefly with describing the diversity of form among insects; discussions on function being correspondingly condensed. In the present work the emphasis is reversed. Struc• ture is described only to an extent sufficient to make the physiological argument intelligible. Every anatomical peculiarity, every ecological specialization, has indeed its physiological counterpart. In that sense, anatomy, physiology and ecology are not separable. But regarded from the standpoint from which the present work is written, the endless modifications that are met with among insects are but illustrations of the general principles of their physiology, which it is the aim of this book to set forth. Completeness in such a work is not possible, or desirable; but an endeavour has been made to illustrate each physiological characteristic by a few concrete examples, and to include sufficient references to guide the student to the more important sources. The physiology of insects is to some the handmaid of Economic Entomology. For although it is not the purpose of physiology to furnish directly the means of controlling insect pests, yet the rational application of measures of control• whether these be insecticides of one sort or another, or artificial interferences with the insect's environment-is often dependent on a knowledge of the physiology of the insect in question. Physiology may thus serve to rationalize existing procedures, or to discover the weak spots in the ecological armour of a species. A knowledge of the ecology of a species is always necessary to its effective control; its ecology can be properly understood only when its physiology is known. 1939 PREFACE TO FOURTH EDITION DESPITE THE turmoil of the intervening years there have been substantial advances in our knowledge of insect physiology since this book was published in 1939. Some of these represent new principles; others illUIninate established notions. In this revised edition an attempt has been made to incorporate this new material into the old framework. It is the aim of such a book as this to provide a framework sufficiently well founded to support the new knowledge but not so rigid as to impede its progress.

v PREFACE TO THE FIFTH EDITION

IN THIS edition a few corrections and alterations have been made in the main text. At the end of each chapter will be found a number of Addenda, which include the most recent advances with cross-references to the pages which they supplement. 1953

PREFACE TO THE SIXTH EDITION DURING THE twelve years that have elapsed since the last revision of this book the study of insect physiology has been transformed. New techniques in electron microscopy, neurophysiology, and many sides of biochemistry have led to spec• tacular advances. The rate at which new material is being published is rising steeply. A complete revision of the text with the incorporation of this new knowledge has been undertaken, and some fifty new illustrations have been introduced. As in earlier editions the bibliography has been restricted-so far as is compatible with providing authority for each statement. That has entailed the addition of more than two thousand new references. It goes without saying, however, that to master the advances that are being made on so many fronts, and to sift the essential discoveries from the non• essential, is becoming beyond the range of a single author. It is only the convic• tion that the physiology of any group of animals must be treated as a unity that has encouraged the present writer to make the attempt.

PREFACE TO THE SEVENTH EDITION

THIS EDITION is a reprint of the sixth edition supplemented by inclusions, amounting to some ten per cent. of the whole text, in which the major advances of the past seven years have been incorporated, together with a few new illus• trations. In order to hold down the cost of reprinting, page references to the new material are given in square brackets at the end of the relevant paragraphs; but all the new entries are printed together, under appropriate captions, in an addenda section at the end of each chapter, where the many new references to the literature will be found.

vi CONTENTS

CHAP. PAGB PREFACE V

DEVELOPMENT IN THE EGG References

II THE INTEGUMENT 27 Properties of the cuticle. Formation and shedding of the cuticle. References III GROWTH 61 Moulting. Metamorphosis. Determination of characters during post• embryonic development. Regeneration. Diapause. References

IV MUSCULAR SYSTEM AND LOCOMOTION Anatomy and histology. Physiological properties of insect muscles. Locomotion. References

V NERVOUS AND ENDOCRINE SYSTEMS Nervous system. Visceral nervous system. Endocrine system. References

VI SENSE ORGANS: VISION 2 15 Compound eye. Simple eyes. References VII SENSE ORGANS: MECHANICAL AND CHEMICAL SENSES Mechanical senses. Hearing. Chemical senses. Temperature and humidity. References

VIII BEHAVIOUR 310 Kinesis and related phenomena. Orientation. Co-ordinated behaviour. References IX RESPIRATION 357 Tracheal system. Development of the tracheal system. Transport of oxygen to the tracheal endings. Elimination of carbon dioxide. Respir• ation of aquatic insects. Respiration of endoparasitic insects. Respir• atory function of the blood. Regulation of respiratory movements. References X THE CIRCULATORY SYSTEM AND ASSOCIATED TISSUES Circulatory system. Haemolymph. Haemocytes. Pericardial cells and so-called 'nephrocytes'. Fat body. Oenocytes. Light-producing organs. References XI DIGESTION AND NUTRITION Fore-gut. Peritrophic membrane. Mid-gut. Hind-gut. Secretions of the alimentary canal. Digestion of some skeletal and other substances of plants and animals. The role of lower organisms in digestion. Nutri• tion. References XII EXCRETION 553 Urine. Intermediary nitrogen metabolism. Malpighian tubes. Histo• physiology of the Malpighian tubes. Accessory functions of Malpighian tubes. Malpighian tubes during moulting and metamorphosis. Cephalic excretory organs and intestinal excretion. Storage excretion. References vii Vlll THE PRINCIPLES OF INSECT PHYSIOLOGY

CHAP. PAGE

XIII METABOLISM 593 Chemical transformations. Some chemical products of insects. Pig• ment metabolism. Respiratory metabolism. References

XIV WATER AND TEMPERATURE Water relations. Temperature relations. References

XV REPRODUCTIVE SYSTEM 700 Female reproductive system. Male reproductive system. Mating, im• pregnation and fertilization. Some factors controlling fertility and fecundity. Special modes of reproduction. Sex determination. Trans• mission of symbiotic micro-organisms. References

INDEX OF AUTHORS

GENERAL INDEX