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Uva-DARE (Digital Academic Repository) UvA-DARE (Digital Academic Repository) The effects of symbiont induced haploid thelytoky on the evolution of brevipalpus mites Groot, T.V.M. Publication date 2006 Document Version Final published version Link to publication Citation for published version (APA): Groot, T. V. M. (2006). The effects of symbiont induced haploid thelytoky on the evolution of brevipalpus mites. General rights It is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an open content license (like Creative Commons). Disclaimer/Complaints regulations If you believe that digital publication of certain material infringes any of your rights or (privacy) interests, please let the Library know, stating your reasons. In case of a legitimate complaint, the Library will make the material inaccessible and/or remove it from the website. Please Ask the Library: https://uba.uva.nl/en/contact, or a letter to: Library of the University of Amsterdam, Secretariat, Singel 425, 1012 WP Amsterdam, The Netherlands. You will be contacted as soon as possible. UvA-DARE is a service provided by the library of the University of Amsterdam (https://dare.uva.nl) Download date:23 Sep 2021 THE EFFECTS OF SYMBIONT INDUCED HAPLOID THELYTOKY ON THE EVOLUTION OF BREVIPALPUS MITES On the cover The front cover contains four pictures of mites representing the four asexual Brevipalpus species studied in this thesis. In clockwise order, starting with the upper picture on the left, the species are: B. californicus , B. phoenicis , uninfected B. obovatus and Cardinium infected B. obovatus . When maintained on common bean ( Phaseolus vulgaris ), cultures of the different species can be distinguished based on the shape and color of the adult females. Female mites of B. californicus and B. phoenicis are more elongated whereas females of the two B. obovatus species are broader. The elongation is most pronounced in B. californicus females which have V- shaped abdomens, whereas the abdomens of B. phoenicis females are more U-shaped. The two B. obovatus species differ mainly in color. The Cardinium infected mites are reddish brown and the uninfected ones are more greenish. The differences between the four species are most clearly visible from older females that have started oviposition already. The back cover shows mites of various life stages in a culture of B. phoenicis . Photos and design: J. van Arkel. The work presented in this thesis was funded by the Netherlands Foundation for the Advancement of Tropical Research (WOTRO), grant number W89- 141, and was carried out at the University of Amsterdam, Institute for Biodiversity and Ecosystem Dynamics. ISBN: 90-76894-67-1 THE EFFECTS OF SYMBIONT INDUCED HAPLOID THELYTOKY ON THE EVOLUTION OF BREVIPALPUS MITES ACADEMISCH PROEFSCHRIFT ter verkrijging van de graad van doctor aan de Universiteit van Amsterdam op gezag van de Rector Magnificus prof. mr. P.F. van der Heijden ten overstaan van een door het College van Promoties ingestelde commissie, in het openbaar te verdedigen in de Aula der Universiteit op donderdag 26 oktober 2006, te 12:00 uur door Thomas Volkert Marie Groot geboren te Alkmaar Promotiecommissie Promotores Prof. dr. S.B.J. Menken Prof. dr. M.W. Sabelis Co-promotor Dr. J.A.J. Breeuwer Overige leden Prof. dr. L.W. Beukeboom Dr. M. Maraun Prof. dr. R. Stouthamer Prof. dr. P.H. van Tienderen Prof. dr. M.K.H. Veith Faculteit der Natuurwetenschappen, Wiskunde en Informatica Contents Samenvatting pp. 1 Summary pp. 9 General Introduction pp. 17 Chapter 1 pp. 31 Cardinium Symbionts Induce Haploid Thelytoky in Most Clones of Three Closely Related Brevipalpus Species Chapter 2 pp. 49 Adaptation in the Asexual False Spider Mite Brevipalpus phoenicis : Evidence for Frozen Niche Variation Chapter 3 pp. 61 Clonal Diversity and Host Plant Specialization in Asexual Brevipalpus Mites Chapter 4 pp. 75 Horizontal Transmission of Cardinium Symbionts Within and Between Closely Related Parthenogenetic Mite Species Chapter 5 pp. 91 Male Function in the Asexual False Spider Mite Brevipalpus phoenicis Chapter 6 pp.103 Recombination and Intragenomic Variation in Asexual Brevipalpus Mites General Discussion pp.125 References pp.133 Dankwoord pp.151 Curriculum vitae pp.154 Samenvatting Theoretische achtergrond Het merendeel van de diersoorten plant zich seksueel voort; om een nieuw individu te produceren moeten een mannelijke en een vrouwelijke gameet fuseren. Het tegengestelde, aseksuele reproductie waarbij vrouwtjes dochters produceren zonder de bijrage van een mannelijke gameet, is zeldzaam. Deze observaties suggereren dat seksuele voortplanting de beste manier van voortplanting is. Aan de andere kant, seksuele voortplanting zou ook minder voordelig kunnen zijn omdat het inefficiënt is en specifieke kosten heeft. Bijvoorbeeld, in seksueel voortplantende soorten produceren de mannetjes niet direct nakomelingen, en verder wordt veel energie besteed aan het zoeken naar een geschikte partner. Deze schijnbare tegenstelling is bekend geworden als de ‘paradox of sex’ en wordt gezien als een van de belangrijkste vraagstukken in de evolutionaire biologie. Een groot aantal theorieën zijn voorgesteld om de paradox of sex te verklaren. Deze theorieën schrijven verschillende voordelen toe aan seksuele voortplanting die de inefficiëntie en kosten ervan compenseren. Op grond van het soort voordeel kunnen deze theorieën in twee groepen worden verdeeld. De theorieën in de eerste groep geven aan dat seksuele voortplanting de aanpassingen en evolutie van soorten versnelt. Hierdoor kunnen seksuele soorten beter omgaan met de veranderingen in hun omgeving. De theorieën in de tweede groep geven aan dat seksuele voortplanting soorten in staat stelt om schadelijke mutaties kwijt te raken. Aseksueel voortplantende soorten ontbreken deze mogelijkheid en het is te verwachten dat schadelijke mutaties in hun genoom ophopen. De theorieën van beide groepen worden ondersteund door wiskundige modellen. Vooral wanneer de verwachte voordelen van verschillende theorieën gecombineerd worden, kunnen de nadelen van seksuele voortplanting volledig gecompenseerd worden. In de laatste jaren zijn er al meer studies gekomen die bewijs voor de theorieën aantonen in levende organismes. Met dit bewijs kan de paradox of sex als verklaard worden beschouwd; seksuele voortplanting is inderdaad voordeliger dan aseksuele voortplanting. Als dat zo is, dan kan men zich terecht afvragen waarom er toch ook nog aseksuele soorten bestaan. Het doel van mijn thesis is het bestaan van een bepaalde groep aseksuele soorten te verklaren: namelijk de aseksuele soorten van het mijten geslacht Brevipalpus . 1 Samenvatting Er zijn drie mogelijke verklaringen waarom aseksuele soorten, tenminste tijdelijk, kunnen voortbestaan. Ten eerste is het mogelijk dat aseksuele soorten speciale aanpassingen hebben die de nadelen van het gebrek aan seksuele reproductie compenseren. Ten tweede, aseksuele soorten zouden kunnen bestaan omdat ze continu worden geproduceerd vanuit seksuele soorten. De nieuwe aseksuele soorten zullen het een relatief korte periode goed doen, maar op langere termijn zullen ze toch extinct gaan. Ten derde, aseksuele soorten zouden kunnen bestaan omdat ze niet volledig aseksueel zijn, maar zich af en toe toch seksueel voortplanten. Niet alleen het bestaan van aseksuele soorten op zich dient verklaard te worden. Een verklaring is ook nodig voor het feit dat veel aseksuele soorten vaak zijn aangepast aan een opmerkelijk brede ecologische niche, een die vaak nog breder is dan de niche van de seksuele voorouder soort. Er zijn twee theoretische modellen die beschrijven hoe een aseksuele soort een brede niche kunnen bezitten. Het ‘Generalist Purpose Genotype’ model voorspelt dat een aseksuele soort zal bestaan uit een klein aantal klonen met zeer brede niches die tolerant ten opzichte van veel verschillende omstandigheden. Dit model gaat er van uit dat specialistische types uitsterven vanwege veranderingen in hun omgeving. Maar, het ‘Frozen Niche Variation’ model voorspelt dat een aseksuele soort zal bestaan uit veel klonen met verschillende smalle niches. Dit model gaat er juist van uit dat generalistische klonen niet kunnen overleven door concurrentie met meer gespecialiseerde klonen. Het model organisme Het genus Brevipalpus is het grootste genus van de familie Tenuipalpidae en telt ongeveer 300 beschreven soorten. Echter, vanwege het kleine formaat van de mijten zijn de taxonomische relaties tussen al die soorten weinig begrepen. Het genus bevat zowel seksueel als aseksueel voortplantende soorten. Deze thesis gaat over de aseksuele soorten. Er is een groot aantal aseksuele Brevipalpus soorten beschreven, maar de drie belangrijkste zijn B. phoenicis , B. obovatus en B. californicus . Het morfologische onderscheid tussen deze soorten is beperkt en intraspeciefieke variatie wordt vaak gemeld. Ieder van de drie soorten komt wereldwijd voor in (sub)tropische gebieden en wordt gevonden op honderden verschillende waardplantsoorten. De meeste waardplantsoorten wordt gedeeld door twee of alledrie de mijten soorten. Deze mijten worden gezien als een belangrijke plaag op groente, fruit en siergewassen. Ze veroorzaken schade zowel direct door te eten van de planten, als indirect door het overdragen van plantvirussen. Bestrijding 2 Samenvatting van de mijten gebeurt doorgaans met chemische middelen. Echter, de mijten ontwikkelen opmerkelijk snel resistenties tegen deze
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