Mechanisms of Self-Incompatibility and Unilateral Incompatibility in Diploid Potato (Solanum Tuberosum L.)

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Mechanisms of Self-Incompatibility and Unilateral Incompatibility in Diploid Potato (Solanum Tuberosum L.) Mechanisms ofself-incompatibilit y and unilateral incompatibility in diploid potato {Solanum tuberosum L.) Promotor: dri rE . Jacobsen Hoogleraar ind e plantenveredeling, inhe tbijzonde r in degenetisch e variatie enreproducti e Co-promotor: drM.S . Ramanna Universitair docent, Departement Plantenveredeling en Gewasbescherming Ronald Eijlande r Mechanisms of self-incompatibility and unilateral incompatibility in diploid potato (Solanum tuberosum L.) Proefschrift ter verkrijging van degraa d van doctor op gezagva n derecto r magnificus van deLandbouwuniversitei t Wageningen dr C.M. Karssen, inhe topenbaa rt everdedige no pmaanda g 14septembe r desnamiddag s te 13.30uu r ind eAul a 'VA/ 3S'V'T Thisthesi s encompasses apar to fth e scientific researchcarrie dou t ondiploi d potato, at the former department ofPlan t Breeding, Wageningen Agricultural University. An international cooperation onself-incompatibilit y (SI)i npotat obetwee npartner si nItal y(Universit yo fSiena) ,German y(Ma x Planck Institute, Cologne) and TheNetherland s (KUN,Nijmege n and WAU, Wageningen) was focused on various aspects of SI. Thisjoin t project was supported by the European Community 'BridgeProgramme ' (BIOT-CT-900172). CIP-DATAKONINKLIJK E BIBLIOTHEEK, DENHAA G Eijlander ,R Mechanisms ofself-incompatibilit y andunilatera l incompatibilityi n diploidpotat o {Solanum tuberosum L.) Thesis Wageningen Agricultural University- with references- with summaries in English and Dutch. Department of Plant Breeding, P.O. Box 386, 6700 AJ, Wageningen,N L ISBN 90-5485-893-1 Cover: seeals oFig.5 ,pag e 66 Key words: Solanum tuberosum, Solanum verrucosum,self-incompatibility , self-compatibility, unilateral incompatibility, unilateral incongruity, 5-glycoprotein, S-RNase, sense, antisense, overexpression. BibliographicAbstract :Thi sthesi sdescribe sth ecreatio nan dselectio no fdiploi d potato genotypes withwel ldefine d self-incompatibility (SI)reactions . Thecontributio n ofth e stylarproduct s ofth e incompatibility alleles, the ^-glycoproteins, is described for both the gametophytic self- incompatibilityreactio nan d theincomplet einterspecifi c crossingbarrie rtha texist sbetwee ndiploi d potato and its self-compatible relative, S.verrucosum. This barrier is called unilateral incompatibility or unilateral incongruity (UI). Complex interactions between incompatibility determining genes are described. BIBLIOTHEEK LANDBOUWUNIVERSITEIT WAGENINGEN Stellingen 1 De aanscherping van de unilaterale incongruentie-hypothese door de S-locus uit te sluiten van de bijdrage aan interspecifieke incompatibilité«(Hogenboom , 1973)i sonterecht . DeU I hypothese heeft in deze vorm derhalve zijn tijd gehad (dit proefschrift). 2 DeRNase-activitei t vanS-glycoproteïne nspeel ti nhe tunilateral e incompatibiliteitssysteem bestaande tussenLycopersiconperuvianum e nL.esculentum een ondergeschikte rol (Rick, 1986; Kowyama et al., 1994; Royo et al., 1994;di t proefschrift). 3 Het "allele-specific dominant negative effect" van een RNase-defecte kopie van een werkzaam incompatibiliteitsallel zoalsbeschreve n door Mc Cubbin et al (1997) iso p zijn best co-dominantie. 4 Alle verwijzingen door de in dit proefschrift genoemde auteurs naar oudere literatuur m.b.t. de uitdrukking "wederzijdse afzwakking (mutual weakening)" betreffen integenstellin g tot wat wordt gesuggereerd niet de uitdrukking maar slechts het verschijnsel. 5 De "two-power competition" hypothese van Abdalla (1970) die beschrijft hoe S.verrucosum en S.tuberosumkruisingstechnisc h van elkaar gescheiden zijn, heeft een hoogantropopatisc h gehalte (= het vertonen van menselijke gevoelens). 6 CytoplasmatischeMannelijk e Steriliteitzoal sdi eoptreed t inhe tsystee m S.verrucosumx S.tuberosum (Abdalla &Hermsen, I97lb) isgee n probleem voor deveredelin g maar een oplossing. 7 Het gebruik van incompatibiliteit ind ehybride-rassen-producti e van zelfcompatibele Solanaceae is een achterhaald idee. 8 Detoekennin g van de soortstatus aanS.sucrense doorHawke s wordt ontkracht door de beschrijving van Hawkes (1989). (Hawkes &Hjertig, The potatoes of Bolivia. 1989). 9 Dedetecti e van 9 (negen!) verschillende incompatibiliteitsallelen in een diploïde Fl van Nicotiana glauca(Pandey , I98l) isee n sterke aanwijzing voor slaperigheid bij de referenten of hoge activiteit van specifieke paramutatie (sic). Dit pleit voor het vermelden van de referenten bij gerefereerde artikelen. 10 Demogelijkhei d vanxenotransplantati e zouvoo r het individu niveau zeer aantrekkelijk zijn, maar ongewenst voor de gemeenschap. 11 Het model zoals dat door de theoretisch natuurkunde Gerard 't Hooft werd gepresenteerd in het programmaNoorderlich t ("Kosmische anarchie",VPRO,28-12-1997 , 20:05 - 20:28) impliceert niet alleen zoals hij zelf alaangaf , predestinatie, maar ook dat de geest een functie isva n de materie. 12 De "Pensee"di e bekend staat als"He t Godsbewijs van Pascal"bewijs t alleen Pascal's opportunisme en zijn feilbaarheid in de kansrekening. 13 Slimheid verhoudt zich tot A.I.O.-schap alsS-homozygoti e tot incompatibiliteit. Stellingen behorende bij hetproefschrif t "Mechanisms of self-incompatibility and unilateral incompatibility in diploid potato (Solanum tuberosumL) " Ronald Eijlander Wageningen, 14septembe r 1998 Voorwoord Ind ejare n zestig enzeventi gwer d eraa nhe t toenmalige Instituut voor Plantenveredeling van de Landbouw Hogeschool Wageningen veel onderzoek gedaan aan aardappel en aan aardappel verwante soorten, endi t gaat in feite tot op dehuidig e dag verder. In die tijd ise r veel bijzonder materiaal ontwikkeld. Een deel van dit materiaal heeft direct of indirect zijn weg gevonden naar aardappelveredelingsbedrijven, terwijl ander materiaal verder gebruikt werd op tal van andere onderzoeksinstellingen.Nakomelinge nva ndri ehee lbijzonder e aarappelplantenvonde nz ohu nwe g naar het Max Planck Instituut te Keulen, alwaar DrRichar d Thompson enmedewerker s opnieuw een aantal eigenaardigheden onder de loupe namen, maar dit keer met modernere, moleculaire technieken. De complicaties in de analyses waren aanleiding om dehul p in te roepen van zowel Prof. Dr Ir Jacobsen's onderzoeksgroep "Genetische variatie en reproductie" van de vakgroep Plantenveredeling als van de emeritus hoogleraar Prof.D rI r Hermsen, die nog steeds actief was op devakgroe p enberei d gevonden werd zijn kennis omtrent dit materiaal met anderen te delen. Dezesamenwerkin gresulteerd euiteindelij k inee nbi jd evakgroe pgeplaatst epositi evoo ree nA.I.O . binnenee nE.E.G . gefinancieerd internationaalproject .Dez epositi ewer ddoo rmi j ingevuld enhie r heb ik dan ook de afgelopenjare n met veelplezie r aan gewerkt. Ditproefschrif t isee nweergav eva nslecht see ndee lva na lhe twer kwa ti ksame nme tvel e anderen aandi taardappelmateriaa l hebverricht .He ti svoo rsommigen ,nie ti nd elaatst eplaat svoo r mijzelf, misschien dan ook frustrerend temoete n zien dat zoveel werk niet inpublicatievor m het daglicht zalzien .D etegenslage n enachtera fonjuis t geblekenwerkhypothese n hebbenmi j enmij n studenten danoo kee nzeker e"faam "opgeleverd . Ikherinne rmi jno gd euitspraa k naar een student toe, toen een experiment wat normaal gesproken nooit mis gaat, maar nujammerlij k de mist in ging, die luidde: "ja , wat hadj e anders verwacht,j e werkt immers bij Ronald". Het heeft deze student er echter niet van weerhouden om ook zelf A.I.O. te worden. Maar het proefschrift is er dan toch gekomen. Ik wil hierbij mijn promotor prof. dr ir. Evert Jacobsen bedanken. Evert, je hebt mij in de gelegenheid gesteld dit tochwe l erg leukeonderzoe k teverrichten . Jeheb t voortdurend de grote lijnen in het oog gehouden en je hebt er voor gezorgd dat e.e.a. hopenlijk toch nog enigszins begrijpelijk oppapie r is gezet. Onshemelsbred e verschil in stijl heeft ervoo r gezorgd datw ehe t nodige geduld met elkaarmoeste nhebben ,maa r wezij n er samentoc huitgekomen . Ikzelfbe n in ieder geval tevreden methe tuiteindelijk e resultaat. Ikhoo p datji j end elezer s dat ook zijn. Verder wil ik mijn co-promotor Dr Ramanna bedanken voor zijn, zoals dat heet, dagelijkse begeleiding. Ik heb u, zoals al vele promovendi voor mij, leren kennen als aimabel, geduldig en onbaatzuchtig. Uvon d altijd tijd ommij n uiteenzettingen aant ehore n envanui t uw grote kennis meet edenken . Ikhe b inu niet alleen eenkundig e collega gevonden, maar ook eenvriend . Prof. Hermsen heb ik reeds in mijn studententijd en ook nu tijdens mijn A.I.O.-schap leren waarderen alsdocen te nal smens .D ehe bme tu vele ,vel eure naa ntafe l doorgebracht, gebogen over papieren vol met afstammingen en genetische modellen, speculerend over wat er nu weer aan de handko nzijn . Uwkenni sva nhe tmateriaa l enhe tverschijnse l "incompatibiliteit" zijn voormi j van onschatbarewaard egeweest .U wbijdrage n aanmij n onderzoek hebben danoo k geresulteerd inee n co-auteurschap, waar umi j een grootplezie rme eheef t gedaan. Dirk-Jan Huigen wil ik bedanken voor al het werk dat hij gemerkt en ongemerkt voor mij heeft verricht.Ji j coördineerde, samenme t demense nva nUnifarm , hetwer k ind ekassen .Toe n iko pd e vakgroep kwam, had jij het voorwerk al gedaan en kon ik zo aan de slag. Je hebt me veel organisatorische takenui thande n genomen enwaarschuwd ewannee r erwee r eenswa thoo i vand e overvolle vork afviel. En tot de promotie aan toe blijf je betrokken bij wat er moet gebeuren. Veertien september iswaarschijnlij k delaatst eda gdatj e nogvoo rm ei n dewee rbent . Bedankt. Marja Schipperse nAnj a Posthuma,julli ehebbe nvoo rmi jhe tnodig ein-vitr
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