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A Multitrophic Approach Promotoren: Prof.Dr .L.E.M .Ve T Hoogleraar Ind Eevolutionair E Ecologie

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A Multitrophic Approach Promotoren: Prof.Dr .L.E.M .Ve T Hoogleraar Ind Eevolutionair E Ecologie Ecology of Drosophila aggregation pheromone: a multitrophic approach Promotoren: Prof.dr .L.E.M .Ve t Hoogleraar ind eevolutionair e ecologie Prof.dr .M .Dick e Hoogleraar ind e insect-plant relaties, inhe tbijzonde r tritrofe interacties Prof.dr .J.C .va nLentere n Hoogleraar ind eentomologi e Samenstelling promotiecommissie: Dr. T.S. Hoffmeister, Christian-Albrechts-Universitat Kiel, Duitsland Prof.dr .S .Daan , Rijksuniversiteit Groningen Prof.dr .R.F .Hoekstra , Wageningen Universiteit Dr. L.Hemerik , Wageningen Universiteit io3>£ Bregje Wertheim Ecology of Drosophila aggregation pheromone: a multitrophic approach Proefschrift ter verkrijging van de graad van doctor op gezagva n de rector magniflcus van Wageningen Universiteit, Prof.dr . ir. L.Speelma n in het openbaar te verdedigen opmaanda g 1 oktober 2001 desnamiddag s te half twee in de Aula. Wertheim, Bregje (2001). Ecology ofDrosophila aggregation pheromone: amultitrophi c approach Thesis Wageningen University, -with ref. -with summary in Dutch ISBN: 90-5808-433-7 Printed byPonse n &Looije n bv,Wageningen , the Netherlands. >d?o\ ,2>o S? STELLINGEN Elk voordeel heb z'n nadeel. J. Cruijff, dit proefschrift Micro-organismen die bij de kolonisatie van nieuwe habitats beperkt worden in hun reproductief succes door lage populatiedichtheden kunnen baat hebben bij het exploiteren van een vector met aggregatiegedrag. Het is waarschijnlijk dat een mutualistische interactie tussen micro-organismen en insecten meerdere malen heeft geleid tot de evolutie van aggregatieferomonen. De veronderstelling dat aggregatieferomonen slechts 'mannelijk geproduceerde sexferomonen' zijn is onaannemelijk. contra Landolt (1997) American Entomologist, 43: 12- 22,di t proefschrift Populatiedynamica wordt wel degelijk in belangrijke mate bei'nvloed door variatie en flexibiliteit in individueel gedrag. contra Ives et al. (1999) The American Naturalist, 154:65 2- 673,di t proefschrift 5. Ieder voedselweb gaat vergezeld van een informatieweb en dat heeft grote gevolgen voor multitrofe interacties. dit proefschrift 6. De kosten-batenbalans van het verbod van de rooms katholieke kerk op condoomgebruik zal negatief uitvallen: het aantal geboorten door het verbod op condoomgebruik zal niet opwegen tegen de sterfte als gevolg van dit verbod. 7. De vrijhandel blijft een farce zolang de rijke landen hun eigen exportmarkt beschermen met exorbitant hoge invoerrechten op alles behalve primaire grondstoffen. Alsd emaatschappi j eenberoe pdoe to pwetenschappelijk e expertise moetnie td emenin g van een wetenschapper de basis vormen voor de maatschappelijke opinievorming maar moet juist de discussie tussen wetenschappers centraal staan, omdat de belangrijkste informatie in de nuances zit. de Avond van Wetenschap en Maatschappij, 6novembe r 2000 9. Het is wrang dat de personeeltekorten bij de overheid voor een belangrijk deel veroorzaakt worden doord e uitstroomva ngemotiveerde ,goe dopgeleid ewerknemer sdi e een gemis ervaren aan toekomstperspectief, terwijl de specifieke overheidstaken - zoals gezondheidszorg, onderwijs en fundamenteel onderzoek - bij uitstek duurzaam zijn. 10. De vergrijzing binnen de Wageningen Universiteit neemt zorgwekkende vormen aan, hetgeen blijkt uit het verschijnen van advertenties van begrafenisondernemers in het universiteitsblad Wb. 11. There is always one more bug. Murphy's software programming law 12. Het gemier over muggen en vliegen draagt bij aan de overlast. 13. Voor wandelvakanties geldt: alles wat je thuislaat is mooi meegenomen. Stellingen behorend bij het proefschrift van Bregje Wertheim 'Ecology of Drosophila aggregation pheromone: a multitrophic approach' Wageningen, 1oktobe r 2001 CONTENTS Abstract Chapter I General introduction and summary 9 Bregje Wertheim Chapter 2 Aggregation pheromones innon-socia l insects:A n ecological and evolutionary 17 perspective Bregje Wertheim, Erik-Jan A. vanBaalen, MarcelDicke &.Louise E.M.Vet Chapter 3 The role of aggregation pheromones in an ecological web:A field study on 45 Drosophila (submitted) BregjeWertheim, RolandAllemand, Louise E.M. Vet &Marce lDicke Chapter4 Behavioural plasticity in support ofa benefi t ofusin g aggregation pheromone in 67 Drosophilamelanogaster (submitted) BregjeWertheim, MarcelDicke &Louise E.M.Ve t Chapter 5 Allee effect in larval resource exploitation inDrosophila :a n interaction 77 between density ofadults ,larva e and micro-organisms (submitted) BregjeWertheim, )ulien Marchais, Louise E.M. Vet &Marce lDick e Chapter 6 Increased risk ofparasitis m asecologica l costs ofusin g aggregation pheromone: 91 Laboratory andfield stud y ofDrosophila - Leptopikna interaction (submitted) Bregje Wertheim, Louise E.M. Vet &Marcel Dicke Chapter 7 Individual risk ofparasitis m in host aggregations: Abehaviou r-base d model on 111 functional and numerical responses (submitted) Bregje Wertheim &Rampa lS . Etienne Chapter 8 The interaction between dispersal, the Allee effect and scramble competition 139 affects population dynamics:a cas e studyo n Drosophila (submitted) RampaiS .Etienne, Bregje Wertheim, Lia Hemerik, Petra Schneider &Ljames Powell Chapter 9 General discussion: Towards an ecological cost-benefit analysis ofaggregatio n 155 pheromone usei nDrosophila melanogaster (submitted) Bregje Wertheim References 163 Nederlandse inleiding en samenvatting 181 Nawoord 191 Curriculum vitae 195 Listo fpublication s 197 ABSTRACT Many insect species use an aggregation pheromone toform group swit hconspecific s in certain localities ofth e environment. This typeo fbehaviou r hasa variet yo fimplication s for ecological interactions, both directly throughth eeffec t ofth epheromon e onth ebehaviou ro fcon - andheterospecifics , andindirectl y throughth econsequentia laggregativ edistribution stha tma yaffec t speciesinteractions .Th e evolutionary ecologyo fth eus eo faggregatio npheromon eha sreceive donl ylittl eattention .Yet ,thes epheromone sma y play an intricate role in food webinteraction s byprovidin g an accompanying information web. The aimo fthi s thesisi st ofurthe r our understanding on the ecological and evolutionary aspects ofth e useo faggregatio n pheromone in insects.B yunravellin g costs andbenefit s that arisefrom th e use ofaggregatio npheromon e inou r ecologicalmode lorganism ,Drosophila melanogaster, westriv et oanswe r why they use an aggregation pheromone and elucidate the ecological consequences of an aggregation pheromone in afoo d web context. In laboratory andfield studies , we identified behaviours and interactions of the fruit fly D.melanogaste r that wereaffecte d byit saggregatio npheromone .Th epheromon e affected thedistributio n ofadults ,thei r eggs, competitor species and parasitoids. Moreover, a number of costs and benefits to the use of aggregationpheromon ewer eindicated .I nsubsequen tstudies ,th emajo r hypotheseso ncost san d benefits were examined. A major benefit of using aggregation pheromone was shown to be aggregated oviposition. Aggregated oviposition enhanced the quality of the larval resource, asindicate d by ahighe r survivalo f the larvae and larger sizeo fth e emergingflies. Thi s Alleeeffec t wascharacterise d bya positiv e effect of adult densityo n larvalfitness components , and mayhav earise nfro m the interaction between adult flies andmicr o-organism s (yeastsan dfungi) .Fung iantagonis eyeas tan dlarva ldevelopment ,whil eadult sca n inoculate yeast on a substrate and temper fungal growth. Larvae also tempered fungal growth, but an increased larval density did not result in an Allee effect but in competition instead. A major cost of using aggregation pheromone arose from an increased risk of parasitism. The parasitoid Leptopilina heterotoma usesth e aggregation pheromone ofadul t fruitflies t o localise the larval hosts,an dbase do nthi sinformatio n thisparasitoi dca ndifferentiat e quantitatively atlon grang ebetwee n substrates that differ inprofitability . After arrivalo n asubstrate , the pheromones nolonge rpla ya rol ei n the host searchingbehaviour . Abehaviour-base d model wasdevelope d topredic t the individual risko f parasitism for hosts indifferentl y sizedhos t aggregations.Th e functional and numerical responses of the parasitoidswer ecombine d witha flexibl e patch leavingdecisio nrul efo r theparasitoid , toasses swhethe r aggregation could also comprise abenefi t to the hosts in terms of adilute d risk (sensu Hamilton 1971). Themode lpredictio nread stha t aggregationi sno tbeneficia l inth econtex t ofth eDrosophila - Leptopilina interaction, and these predictions were supported byfield data . Ina simpl espatio-tempora l simulationmodel ,th epopulatio ndynamic sarisin gfrom severa lmode s ofdispersal ,foo d competition and anAlle e effect wereexplored .Th e modeli sa first ste ptoward sa mor e extensive model that incorporates the responses of insects to spatially heterogeneous resources and chemical information (e.g.,aggregatio n pheromone). The main conclusion from this thesis is that the aggregation pheromone of D. melanogaster plays an intricate role within afoodwe b context, and that avariet y ofcost s and benefits arisefrom multitrophi c interactions. To understand the dynamic interactions in this and many other ecological systems, it is essential togai n more insight into the effect ofaggregatio n pheromone on the behaviour ofindividuals . CHAPTER 1 General introduction and summary Evolutionary ecology The distribution of individuals across the environment islargel y determined bybioti c interactions and environmental characteristics
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