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In the Chrysomelid Beetle,Galeruca Tanaceti (Linn.)

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In the Chrysomelid Beetle,Galeruca Tanaceti (Linn.) SOME PHYSIOLOGICAL ASPECTS OF ADULT REPRODUCTIVE DIAPAUSE IN THE CHRYSOMELID BEETLE,GALERUCA TANACETI (LINN.) by Yow Cheong, Siew, M.Sc. (1.Z.) Thesis submitted for the Degree of Doctor of Philosophy in the Faculty of Science, University of London Imperial College of Science and Technology, Field Station, Silwood Park, Sunninghill, Ascot, Berkshire. May, 1963. ABSTRACT Galeruca tanaceti Linnaeus is a short-day insect. Photoperiod and temperature are the primary causal factors which induce, sustain and terminate diapause. The critical photoperiod is about 13 hours at 20°C, under controlled conditions, whereas in the field it is about 14.5 hours, where the daily mean air temperature fluctuates around 15°C. Diapause can be sustained, terminated or reinstated depending on the photoperiod to which the beetle is subjected at any period of its adult life. The phases of diapause, ovarial maturation and oviposition are controlled by different levels of hormonal activity. Diapause is sustained by a low level of activity of the neurosecretory cells of the brain, which in turn influence a low level of activity of the neurosecretory system. Five types of neurosecretory cells, (A, A1, B, C and D) occur in the brain and the suboesophageal ganglion. These cells undergo short cycles of secretory activity throughout adult life. These cycles become shorter but more intense towards and at the phase of oviposition. The results stem from two experimental sources: (1) measurements of nuclear volumes of these cells and (2) autoradiographic investigations using 35S-DL-cystine. The corpus cardiacum/allatum complex functions in relation to the level of activity in the neurosecretory cells. Three criteria were used to determine the corpus cardiacum/allatum activity: (1) increase in volume of the glands, (2) decrease in nuclear-cytoplasmic ratios and (3) increase in nuclear volumes. To elucidate the role of the components of the neurosecretory system, cauterization and implantation experiments were carried out. These experi- ments indicate a close inter-relationship between the neuro-endocrine centres and the possibility of complex feed-back mechanisms. A comparison is made of the respiratory rates, fat and water contents in the pre-diapause, diapause and ovarial maturation phases. ii. TABLE OF CONTENTS Page I. Introduction a) General Ooo 04. 0.0 ••• •o• ••• ••• ••• 1 b) Review of the literature ••• ••• ••• ••• ••• 3 II. The Life History of G. tanaceti ••• ••• ••• ••• ••• 17 III.Rearing techniques ••• ••• ••• ••• ••• ••• ••• 19 IV. Studies on the morphology of the reproductive systems with accompanying quantitative and qualitative changes at different stages of the adult life of G. tanaceti Introduction . • • • • • ••• ..• ••• ••• ••• ••• 20 Method • ee ... ... ••• ... do• • A • • ... ... 20 a) Morphology of the reproductive organs ... 000 004 21 b) Quantitative and qualitative changes of the reproductive organs at different stages of adult life history 604 23 Discussion ••• ••• SO• ••• 26 V. The neuro-endocrine complex of G. tanaceti Introduction 004 4.0 00. 4.0 6.0 006 60.. ego 29 Method ... 000 0.0 0.0 006 600 000 000 000 30 Morphology and anatomy of the neuro-endocrine complex 31 The histology of the neurosecretory system 4104. 00. se. 35 Discussion • • . • • • • • • • • • • • • • • • •• ••• 43 VI. Studies on feeding, growth, respiration, water and fat content of G. tanaceti at different stages of its adult life history Introduction • . • • . • • • • • • • • • • • • • • • • 46 1. Feeding and growth 000 0.. 6.0 ••• 47 2. Respiration • .• • • • • • • • • • ••• 49 3. Water and fat contents • • • • • • • • • ••• 54 Discussion • • • • • • • • • • • • • • • • 0 • 59 VII.Experiments on the effects of photoperiodism and temperature on the induction and termination of reproductive diapause and on growth of G. tanaceti Introduction 000 600 00. .00 4.9 6.6 *elk 63 Methods ••• ••• ••• ••• ••• ••• ••• ••• ••• 64 Results ••• ••• ••• ••• ••• • •• ••• ••• ••• 65 Page Rate of maturation and oviposition of G. tanaceti kept at different pht-toperiods and temperatures ... 000 ... 68 Fecundity of G. tanaceti kept at different photoperiods and 4641 ..0 72 temperatures ... ... .00 0.• 0.0 000 Ovarial maturation of G. tanaceti kept at different photo- periods and temperatures ... 000 000 000 000 040 72 The sensitive stage of G. tanaceti in response to photoperiod- ism and temperature 0.0 000 000 40. 000 0.0 78 Growth of reproductive organs at the photoperiod of 12 hours at 2000. e... ... ... ... deo ... ... 4,.. 78 Reversible effects of photoperiodism and of temperature in G.:tanaceti 4,01. 060 0.0 004 0.0 41.. 00. 000 81 Discussion O.. .00 Ote $8. 610. 060 0.0 06. 81 VIII. Studies on the cyclical changes of the neurosecretory system during adult life history of G. tanaceti Introduction 'OS 00. ... 0.0 04.0 see el. ... 85 Methods 004 0.6 000 066 Ott 4..6 0100 85 R esults ... 0.4 .00 000 4,00 000 1.0. 400 60* 89 1. Changes in the N.S.C. of the pars intercerebralis 89 2. Changes in the N.S.C. of the lateral groups of the brain 95 3. Changes in the N.S.C. of the suboesophageal ganglion 98 4. Qualitative and quantitative changes in the corpora cardiaca op .00 800 000 00. Ott 06. 0.4 103 5. Qualitative and quantitative changes in the corpora allata 0•• ••• ••• ••• ••• •O• ••• ••• 107 Discussion • •• •• • • •• • • • •• • ••• •• • • 120 IX. The dynamics of neurosecretion Introduction • • •• .• • • •• .•• • •• ••• •• • 124 Methods > 0.6 000 SOO 000 Ott .00 0.0 .00 125 Results 000 O.. .0. 600 660 64,0 600 Ott 127 a) Demonstration of short range secretory cycles in the N.S.C. by the use of 3 S-DL-cystine in auto- radiographic study . ... ... eel e.. *of 127 b) Experimental demonstration of a possible "neuro- endocrine momentum" 00. $00 0010 Ott 4100 132 Discussion ... 000 000 .0. 000 00. .00 000 135 iv. Page X. Experimental investigations into the role of the neurosecretory system in controlling reproductive diapause, ovarial maturation and oviposition in G. tanaceti Introduction .• • • •• •• • • a • • • • • • • II • • • • • • 141 Methods •• • ... •• • •• • ... •• • • •• •• • • • . 141 Results ... • • • ... •• • •• • •• • ... ... ... 144 a) Cauterization experiments ... •• • •• • •• • •• • 144 b) Implantation experiments ... •• • •• • ... •• • 150 Discussion . •• • •• • •• • ••• •• • ••• ... •• • 152 XI. General discussion • • • • • ... •• • •• • ... •• • • • • 156 Summary .. • • • III • • • • • • • • • • a • • • • • • • • 0 • • • • 168 Acknowledgements ... •• • •• • ••• •• • ... •• • ... • • • 171 Bibliography .. • •• • • • • • • •• • •• • ••. •• • ... ... 172 1. I. INTRODUCTION a) General The phenomenon of diapause was first demonstrated by Duclaux (1869). He observed that the egg batches of Bombyx mori invariably failed to hatch at room temperatures whereas, similar batches which had been chilled previously for 40 days in an ice box, hatched successfully when returned to the higher temperature. This phenomenon of diapause in insects has recently stimulated much research as is evident from the reviews by Bonnemaison (1945), Andrewartha (1952), Lees (1955, 1956), Church (1955), Hinton (1957), Willer (1957a), Danilevskii (1961), Harvey (1962), de Wilde (1962) and van der Kloot (1960, 1962). The problem of arrested growth has become appreciated as a highly complex mechanism for the preservation of the species in regions where seasonal climatic conditions are unfavourable for continuous growth and for reproduction. It has long been recognised that the environment ultimately controls diapause. During the last 25 years, however, with the classical work of Kogure (1933) on Bombyx mori, investigations into the effects of photoperiodism and temperature have shown that these are the primary causal factors, inducing, sustaining and terminating diapause. From the physiological standpoint, these external factors form only part of the problem. The other aspect of the problem revolves mainly around the physiological and bio-chemical control of growth. It is now known from the classical research of Wigglesworth, Fukuda and Williams that the cessation of growth and the accompanying metabolic adjustments are under hormonal control. The neurosecretory system which has been under active investigations during the last 15 years, contains 2. the main endocrine centres. The neurosecretory system is thought to provide the link, coordinating the internal physiology of the insect with the external environment. Indeed, it is this link, as Lees (1955) suggests, that permits the mechanism underlying diapause to function as a timing device, synchronizing the periods of dormancy and active growth with the seasonal changes in the environment. There has not been any attempt so far by workers of diapause to investigate the physiology of imaginal diapause as outlined in the above scheme. de Wilde and his associates (1958, 1959, 1960) partially approached the problem in their study on the role of the corpus allatum as an endocrine centre, controlling imaginal diapause. The principal theme in this study is to consider the role of the neurosecretory system in the control of induction and termination of diapause, in response to the effects of photoperiodism and temperature. The Chrysomelid beetle, Galeruca tanaceti L., passes through well- defined physiological phases in its adult life. Because of this, it offers possibilities of an integrated investigation into the external factors controlling diapause and the internal physiology of the insect. I have divided the adult life history into four physiological phases,
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