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^Ùcj\( Promotor: Dr Selection of Trichogramma for inundative biological control A study of behavioural variations among strains and species of an egg-parasite genus CENTRALE LANDBO UW CATALO GU B 0000 0262 5396 ^ùCj\( Promotor: dr. J. C. van Lenteren hoogleraar in de Entomologie, in het bijzonder de oecologie der insekten Gerrit Adriaan Pak Selection of Trichogramma for inundative biological control Proefschrift ter verkrijging van de graad van doctor in de landbouwwetenschappen, op gezag van de rector magnificus, dr. C. C. Oosterlee, in het openbaar te verdedigen op woensdag 11me i 1988 des namiddags te vier uur in de aula van de Landbouwuniversiteit te Wageningen BIBLIOTHEEK LANDBOUWUNIVERSITEIT WAGENINGEN /S/V* 2J-$fé>< "The behaviour of insect parasites plays a great part in fixing theirvalu e aspes t controls;an d itfollow s that we must know their behaviour before we can use them efficiently." G. Salt,195 8 Copyright © 1988 by Gerrit A. Pak All rights reserved Omslag: P.J. Kostense Druk:Grafisc h bedrijf Ponsen enLooije n - Wageningen ISBN 90-9002176-0 /O^o^o', itZù Stellingen 1. Het mislukken van de toepassing van Trichogramma voor de bestrijding van insektenpla- gen is in de meeste gevallen het gevolg van onvoldoende kennis van inter- en intraspecifieke variatie in het zoek- en parasiteringsgedrag van vrouwelijke wespen binnen dit genus. F. D. Parker & R. E. Pinnell. Environmental Entomology 3: 935-938 (1974). Dit proefschrift. 2. Trichogramma vrouwtjes zijn, door beoordeling van de kwaliteit van potentiële gastheren, in staat hun eileggedrag op adaptieve wijze aan te passen aan hun gastheer. G. Salt. Proceedings of the Royal Society London (B) 122: 57-75 (1937). M. J. van Dijken &J . K. Waage. Entomologia Experimentalis et Applicata 43: 183-192(1987).- Dit proefschrift. 3. De hypothese van Klomp et al. dat Trichogramma vrouwtjes het vermogen tot gastheer­ discriminatie moeten leren door eilegervaring met een ongeparasiteerde gastheer, en dit vervolgens kunnen vergeten, gaat voorbij aan het belang van idee-vorming over wat naïeve wespen feitelijk door een dergelijke ervaring geleerd zouden kunnen hebben. H. Klomp, B. J. Teerink & W. C. Ma. Netherlands Journal of Zoology 30: 254-277 (1980). J. E. R. Staddon. Adaptive behavior and learning. Cambridge University Press, Cambridge (1983). 4. Het veldonderzoek van Kareiva naar de invloed van habitat opdeling op de stabiliteit van een predator-prooi systeem toont het belang aan van het toetsen van hypothesen over het gedrag van predatoren en parasieten in natuurlijke gemeenschappen van hun prooien of gastheren. P. Kareiva. Nature 326: 388-390 (1987). 5. Het primaire doel van insektenbestrijding is niet de bescherming van gewassen, maar het beschermen van landbouwbedrijven tegen een mogelijk faillissement. J. H. Perkins. Insects, experts, and the insecticide crisis. Plenum, New York (1982). 6. Bestrijding van de illegale teelt van coca planten in Peru met behulp van de vlinder Eloria noyesi zou kunnen leiden tot een entomologische wapenwedloop. Butterflies thwart cocaine barons. New Scientist 1598: 27 (4-2-1988). 7. Betiteling van de meest effectieve sluipwesp voor biologische bestrijding van Insektenpla­ gen als „Porsche onder de sluipwespen", kan de onterechte indruk wekken dat hier sprake is van een relatief dure methode van bestrijding. De Porsche onder de sluipwespen - biologische bestrijding van mineervliegen. Wagenings Universiteitsblad 31:8 (15-10-1987). 8. De bedrijfsmatige productie van natuurlijke vijanden voor biologische bestrijding kan bevorderd worden door de invoering van „kwekersrecht" voor de geselecteerde en in massakweek gehouden stam van de natuurlijke vijand. 9. Intelligentie blijkt niet alleen uit het verstandelijk vermogen om dingen te doen, maar evenzeer uit het vermogen om dingen te laten. R. E. Leaky &R . Lewin. Origins. Dutton, New York(1977) . 10. Het feit dat wetenschappelijk onderzoek intoenemend e mate gericht iso p het oplossen van problemen veroorzaakt door toepassingen van onderzoek, zou tot diepgaande bezinning moeten leiden. 11. Discriminatie van vrouwen op de arbeidsmarkt zal verminderen indien mannen in gelijke mate als vrouwen hun loopbaan aanpassen aan die van hun partner en aan eventuele kinderzorg. Vrouwen op de arbeidsmarkt - de aandelen stijgen. Onderzoek NILI-MPW, Wageningen (1987). 12. Ministers zouden net zo gemakkelijk afstand moeten doen van hun functie als van hun dienstauto. 13. Advertenties met deteks t „roken moet mogen"zij n eenbeledigin g voor mensen died e rook niet willen zien of kunnen luchten. 14. Bij de pakken neerzitten kan een aangename bezigheid zijn. Stellingen behorende bij het proefschrift van G. A. Pak: „Selection of Trichogramma for inundative biological control". Wageningen, 11 mei 1988. Panorama Wie door dit doek heen stapt verkijkt zich op demaa t van zijn bereik.D e wereld is maar rond ennau w om ons gesteld. Vanuit een punt gezien eenweid s gezicht, met mogelijkheid tot wisseling van zijn; alleen beweging biedt het juist perspectief: er komt een eind aanwa t geen eind wou zijn. T.va n Deel Achter de waterval Gedichten Querido,198 6 VOORWOORD Volgens het promotiereglement vand eLandbouwuniversitei t dient een proefschrift "debekwaamhei d vand e promovendus tot de zelfstandige beoefening van de wetenschap aan te tonen".Toe n ik in 1981 met mijn promotieonderzoek begon,ha d ikweini g weet van de betrekkelijkheid van deze eis.Bi jhe t schrijven van de laatste zinnen van mijn proefschrift word ik stilva nhe t bewustzijn dat dit werkstuk het resultaat isva n de bereidwilligheid vanvel emense n om met mijme e tewerken ,i nhe t bijzonder medewerkers vand evakgroe pEntomologi e enva n de Gecombineerde Diensten Binnenhaven van de Landbouwuniversiteit Wageningen.Ee n ieder die,gevraag d ofongevraagd , bewust ofmisschie n welonbewust ,o pd eêé no f andere wijze heeft bijgedragen aand e totstandkoming vandi t proefschrift,wi l ikdaaro m heel hartelijk bedanken.Zonde r jullie hulp zou 11 mei 1988 voor mij geen feestdag kunnen zijn! Een aantal personen verdient het om opdez e plaatsme t name genoemd te worden,hoewe li khoo phe n reeds eerder blijk gegeven te hebben van mijn waardering.Mee r dan wie ook geldt dit mijn vrouw Els.Veeleisen d als ik ben heb ik het uiterste van jegeverg d aan medewerking,gedul d en liefde, vooral inhe t afgelopen jaar,waari n jeoo kno gkampt eme t een slepende ziekte.He t feit dat onze relatie zich ontwikkelde terwijl je een gedeelte van dit onderzoek verrichtte,vergroo t voor mijd e waardeva ndi t proefschrift. Naast Els de Jong heb ik diverse andere studenten begeleid bijhu n leeronderzoek aan Trichogramma.I ndi t proefschrift zijn resultaten opgenomen van experimenten, die inchronologisch e volgorde verricht werden door Tanja vanHeiningen ,Ingri d Heck,Moniqu e Hermans,Harrie t Buis, Anneke Kaskens, Astrid Nieland,Ro bLindeboom , Frans vanAlebeek ,Kari n Driessen,To n van Dalen,Lamber t Ebbers,Ruu d deGunst ,Carolin e Kersten en Nanda Kaashoek.Al s gastmedewerkers waren bijhe t onderzoek betrokkenH a Quang Hung,Doek e van der Schaaf en Gerti Horeman.Gedurend e twee seizoenen begeleidde Frans vanAlebee k op creatieve ennauwgezett e wijze het veldonderzoek. Gastheerinsekten waren dankzijhe tvakkundi g kweekwerk vanLe oKoopman , Frans vanAggele n en Richard Pieters meestal in overvloed voorradig. Bea Heufkens, Gerard Pesch enHerma n Dijkman verzorgden in de loopde r tijdd e Trichogramms kweek en verleenden assistentie bijdivers e proeven. Bovendien waren Herman's hand- en spandiensten een onmisbare hulp bijd e uiteindelijke vervaardiging van het proefschrift. De technische afwerking van een proefschrift is afhankelijk van de kunde en het enthousiasme van diverse specialisten.Maria n Koopman,Truu s deVries ,Wille m Westland envoora lHannek e Westra werden moe,maa r niet moedeloos,va nhe t welhaast eindeloze typewerk.Voo r de figuren tekenden Wim Middelplaats,Frederi k von Planta,Ste f Wever en bovenal Piet Kostense. Piet ben ik zeer erkentelijk voor de fraaie tekeningen enhe t schitterende omslag.D e mannen van de fotografische afdeling, in het bijzonder de energieke Jan Bakker,maa r ookJac k Brangert,Berr y Geerligs,Wi m van Hof enHei n Visser,gave n met hun dienstverlening inhoud aan de "Gecombineerde Diensten". Voor het schrijfwerk heb ik me gedurende eenreek sva nmaande n zovee l mogelijk afgezonderd. Ikdan k mijn collega'sva n de vakgroep voor hun begrip voor deze situatie.Fran s van Alebeek, Elsd e Jong, Oscar Minkenberg,Louis e Vet enJonatha n Schmidt ben ik bovendien erkentelijk voor hun commentaar op delen van de tekst en suggesties voor verbetering. Bijhe tverkrijge nva nee n promotie-assistentschap waren de inspanningenva nd evroeger evakgroep-medewerker s Gert Ankersmit en wijlen Jan de Wilde van groot belang.Ee n promovendus is bovenal niets zonder een promotor. Als zodanig heeft Joop vanLentere n een centrale rol gespeeld bij het totstandkomen van dit werkstuk.I k ben je vooral dankbaar voor het in mijgesteld e vertrouwen en jevoortdurend e inspiratie.Bovendie n gaf je me debenodigd e ruimte voor een zelfstandige beoefening van de wetenschap, waarvan dit proefschrift de bekroning is. Tenslotte een woord vandan kaa nmij n ouders,omda t jullie me de vrijheid gaven om me teontplooie n ind evoo r jullievreemd e wereld van wetenschap en onderzoek. Gé Pak maart 1988 CONTENTS Curriculum vitae (inDutch ) 11 Publications 12 Summary (inDutch ) 15 Outline of the thesis 19 1. GENERAL INTRODUCTION Biological control 21 Trichogramma 22 Lepidopterous pests on cabbage 24 2. SELECTION OF CANDIDATE NATURAL ENEMIES Pre-introductory research 29 Criteria for selection 29 Selection of Trichogramma 31 Inter- and intraspecific variation 32 Research program 34 3. GENERAL MATERIALS AND METHODS Parasite cultures 37 Host cultures 38 Observations 39 4. ADAPTABILITY TO FIELD-TEMPERATURE CONDITIONS Abstract 43 Introduction 43 Materials and methods 45 Results 48 Discussion 54 5. HOST-AGE SELECTION Abstract 57 Introduction 58 Review of the literature 59 Materials and methods 65 Results 68 Discussion 76 6. HOST-SPECIESSELECTIO N Abstract 81 Introduction 81 Materialsan dmethod s 84 Results 88 Discussion 101 7.
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