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Dynamics of Feeding Responses in Pieris Brassicae Linn. As a Function

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Dynamics of Feeding Responses in Pieris Brassicae Linn. As a Function 595.789:591.185.31:591.513:591.53 DYNAMICS OF FEEDING RESPONSES IN PIERIS BRASSICAE LINN AS A FUNCTION OF CHEMOSENSORY INPUT: A BEHAVIOURAL, ULTRASTRUCTURAL AND ELECTROPHYSIOLOGICAL STUDY (WITH A SUMMARY IN DUTCH) PROEFSCHRIFT TER VERKRIJGING VAN DE GRAAD VAN DOCTOR IN DE LANDBOUWWETENSCHAPPEN OP GEZAG VAN DE RECTOR MAGNIFICUS, MR. M. POLAK, HOOGLERAAR IN DE RECHTS- EN STAATSWETENSCHAPPEN VAN DE WESTERSE GEBIEDEN, IN HET OPENBAAR TE VERDEDIGEN IN DE AULA VAN DE LANDBOUWHOGESCHOOL TE WAGENINGEN OP WOENSDAG 14 JUNI 1972 TE 16.00 UUR DOOR MA WEI CHUN H. VEENMAN &ZONE N N.V. - WAGENINGEN - 1972 Dit proefschrift met stellingen van MA WEI CHUN landbouwkundig ingenieur, geboren te Amsterdam op 17augustu s 1941,i s goedgekeurd door de promotor, DR. J. DE WILDE, hoogleraar in het dierkundig deel van de Plantenziektenkunde. De Rector Magnificus van de Landbouwhogeschool, J. M. POLAK Wageningen, 28 april 1972 Thisthesi s isals opublishe d as Mededelingen Landbouwhogeschool Wageningen 72-11(1972 ) (Communications Agricultural University Wageningen, The Netherlands) STELLINGEN I Bij pogingen tot kausale analyse van elektrofysiologische activiteit in indivi- duele zintuigcellen en daaropvolgend gedrag dient in sterke mate rekening te worden gehouden met de aard van bij de gedragsanalyse betrokken responsie- parameters en experimentele omstandigheden. Dit proefschrift II In de gangbare terminologie bij de klassifikatie van chemicalien met betrek- king tot hun gedragseffekt zouden stoffen met een vraatremmende werking of 'feeding inhibitors' aan de hand van hun werkingsmechanisme kunnen worden gedifferentieerd in 'feeding deterrents' en 'anti-stimulants'. Dit proefschrift III Tumoren bij insekten zijn in demeest egevalle n geen neoplasma's. IV Nederland kan een belangrijke bijdrage leveren aan een kwalitatieveverbete - ring van het dieet in ontwikkelingslanden door verbeterde cultuurmaatregelen, veredelings- en verwerkingsprogramma's van tropische en subtropische peul- vruchten te bevorderen. V De volgens TAKEMOTO et al. in moerbeiblad voorkomende hoeveelheden fyto-ecdysonen zijn te laag in concentratie om van enigerlei betekenis te zijn in het voedselopnamegedrag van dezijderup s Bombyxmori L. Dezelfde stoffen zouden na orale opname welee n invloed kunnen uitoefenen opd e fysiologische ontwikkeling. TAKEMOTO, T., S. OGAWA, N. NISHIMOTO, H. HIRAVAMA, S. TAMIGUCHI, 1967. Yakugaku Zasshi, 57(6):748 . VI Debetekeni sva n apterebladluize n bij deverspreidin g van een non-persistent virus binnen een perceel wordt vaak overschat. VII In het huidige onderzoek over insektenchemosterilantia dient uit oogpunt van praktische toepasbaarheid meer aandacht te worden geschonken aan in- sektenhormonen en hormoonmimetica. VIII Voor vele sekundaire plantestoffen geldt dat zij primair een ecologische functie bezitten. IX De suggestieva n ELIZAROV etal . datcontac t chemoreceptoren ind e mandi- bulae van Agriotes Iarven aanwezig zouden zijn wordt niet door voldoende onderzoek gestaafd. ELIZAROV, J. A., E. S. ZOLOTAREV, T. S. ZULEGOVA, 1968. Vestnik Moskov. Univ., 6: 17-21. X Een beperking ofverbo d van deprodukti e van DDT ind eontwikkeld e lan- den zal niet kunnen geschieden zonder zeer ernstige nadelige konsekwenties voor de ontwikkelingslanden. Proefschrift MA WEI CHUN Wageningen, 14juni197 2 Aanmijn ouders Aan Yuet Wah SueWen CONTENTS 1. GENERAL INTRODUCTION 1 2. MATERIALS AND REARING METHODS 3 2.1. Experimental animals 3 2.2. Plant material 4 2.3. Chemicals 4 3. ASPECTS OF LARVAL PREFERENCE BEHAVIOUR 5 3.1. Introduction 5 3.2. Materials and methods 5 3.3. Experience-dependent modification of preference behaviour . 6 3.4. The effect of mouth part ablation on preference behaviour 14 3.5. Discussion 17 4. QUANTITATIVE STUDY OF BEHAVIOURAL CHEMORESPONSES ... 19 4.1. Introduction 19 4.2. Materials and general methods 19 4.3. Results and discussions 20 4.3.1. Observations on behaviour patterns 21 4.3.2. Parameters of larval chemoresponse 23 4.3.3. Behaviour responses to sugars and related compounds 25 4.3.4. The effect of starvation on the response to sucrose . 29 4.3.5. Behaviour responses to sinigrin and its interaction with sucrose 32 4.3.6. Stimulating effects of some other chemical compounds 36 4.3.7. The effect of calcium and sodium chloride on response to sucrose .... 40 4.3.8. The effect of mouth part ablations 42 4.3.9. Responses of maxillectomized larvae to sugars 44 4.3.10. The effect of mouth part ablation on the response to sinigrin 45 4.3.11. Inhibition of food intake by specific stimuli 48 5. MORPHOLOGICAL AND ANATOMICAL OBSERVATIONS ON SENSORY ORGANS 59 5.1. Introduction 59 5.2. Materials and methods 60 5.3. The maxillary sensilla styloconica 60 5.4. Observations on epipharyngeal sensilla 74 5.5. The sensory structures associated with mandibular fine canals 81 5.5.1. Introduction . 81 5.5.2. Materials and methods 82 5.5.3. Results and discussions 82 6. ELECTROPHYSIOLOGICAL STUDIES ON CHEMORECEPTOR RESPONSES 90 6.1. Introduction 90 6.2. Materials and methods 90 6.3. Results and discussions 91 6.3.1. Ss I response to stimulation by carbohydrates 91 6.3.2. Ss II responses to stimulation by carbohydrates 95 6.3.3. Interspecies specificity towards various sugars 95 6.3.4. Ss I chemoreceptor cell sensitive to specific feeding inhibitory compounds . 98 6.3.5. Cellular specificity 98 6.3.6. Electrophysiological properties 102 6.3.7. Discussion 107 6.4. Electrophysiological studies on the contact chemosensory function of the epipharyngeal papilla-like organs 108 6.4.1. Ep responses to stimulation by some electrolytes 109 6.4.2. Ep responses to stimulation by carbohydrates 113 6.4.3. The effect of sodium chloride on the response of the Ep sugar receptor . 116 6.4.4. The effect of calcium chloride on the response of the Ep sugar receptor . 118 6.4.5. Theoretical considerations on sugar receptor stimulation in SsI I and Ep sugar receptors 119 6.4.6. Responses of Ep to stimulation by specific feeding inhibitors ....... 124 6.4.7. Additional remarks on the Ep innervation 125 6.5. Summary 127 7. THE RELATION BETWEEN BEHAVIOURAL AND PERIPHERAL NEURAL RESPONSES 128 7.1. General considerations 128 7.2. Observations on feeding behaviour 129 7.3. Sensory control mechanisms in feeding behaviour 130 7.4. Mechanisms in inhibition of feeding responses 136 7.5. Final remarks 139 8. SUMMARY 142 9. ACKNOWLEDGEMENTS 146 10. SAMENVATTING 147 APPENDICES 151 REFERENCES 157 1. GENERAL INTRODUCTION Chemoreception plays a crucial role in the interaction of organisms with their environment. In animals and somemicro-organism s chemoreception iso f paramount importance not only in securing food necessary for the subsistence of the individual organism but also in social and other activities necessary for themaintenanc eo fth especies .Th epresen twor k dealswit h chemoreception in relation to feeding and food selection behaviour of larvae ofPieris brassicae (Lepidoptera: Pieridae). Apart from mechanisms based on host avoidance (repellence), the determination of food specificity in phytophagous and other animalsi scause d bydiscriminatio n duringcontac t ofth esubstrat ewit hth eleg s or mouth parts. One of the first to demonstrate experimentally the perception of plant chemicals by phytophagous insects was VERSCHAFFELT (1910) who noticed that food selection by the larvae of P. brassicae and P. rapae was in­ fluenced byth e presence of a distinct class of chemicals (glucosides of allyliso- thiocyanates) in the natural hostplants (Cruciferae, Tropaeolaceae or Reseda- ceae). Since then numerous reports have appeared on chemoresponses, mainly of phytophagous insects as fostered by the obvious economic relevance (for extensivereview s see BECK, 1965; DETHIER, 1947, 1966; SCHOONHOVEN, 1968). However, besides the thorough studies of tarsal and labellar chemoreceptors ofblowflie s (reviewed by DETHIER, 1969),insec tfeedin g behaviour has onlyi na few cases been studied in relation to electrophysiological analysis of chemore- ceptive phenomena. With regard to phytophagous insects such studies are restricted to some lepidopterous larvae (for reviews see SCHOONHOVEN, 1968; DETHIER, 1970) and two coleopterous species,viz .th e Colorado beetleLeptino- tarsadecemlineata (STURCKOW and QUADBECK, 1958, STURCKOW, 1959) and Chrysomela brunsvicensis (REES, 1969). All these studies are more of a qualita­ tive than of a quantitative nature. The lack of more quantitative studies may seem understandable in view of the major difficulties which are related to an attempt to establish a meaningful correlation between quantitative behavioural responses and electrophysiological phenomena in sense organs. One of these problems iscause d by the fact that feeding preferences may be altered through experience or learning. As will appear in Chapter 3 feeding preferences in the larvae of P. brassicae are modifiable within certain limits. Another problem arises from the possibility that the outcome of behavioural studies may vary according to the response parameter chosen. These and other difficulties pose special problems as to the choice of experimental designs and response para­ meters which can be profitably applied to assess larval chemoresponses in a quantitative manner. These issues are dealt with in Chapter 4. In Chapter 5 morphological and anatomical studies of sense organs involved in feeding are described. It will be clear that only an understanding of the organization and function of thecomplet e chemoreceptive system willallo w a quantitative inter­ pretation of the observed feeding responses in relation
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