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Physiological Functions of Phytochromes in Tomato: a Study Using Photomorphogenic Mutants

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Physiological Functions of Phytochromes in Tomato: a Study Using Photomorphogenic Mutants Physiologicalfunction so fphytochrome si ntomato : astud yusin gphotomorphogeni c mutants Fysiologische functies van fytochromen in tomaat: een studie gebruikmakend van fotomorfogenetische mutanten Promotor: dr.W.J . Vredenberg hoogleraar in de plantenfysiologie metbijzonder e aandacht voor de fysische aspecten Co-promotor: dr. R.E. Kendrick universitair hoofddocent bij devakgroe p Plantenfysiologie pt,^oM\^, LeonardusHubertu s Joseph Kerckhoffs Physiologicalfunction so fphytochrome si ntomato : a study using photomorphogenic mutants Proefschrift ter verkrijging van de graad van doctor op gezag van derecto r magnificus van de Landbouwuniversiteit Wageningen, dr. CM. Karssen, in het openbaar te verdedigen op vrijdag 20 december 1996 des namiddags te vier uur in deAula . isn 31,11(02 This research was supported by a grant from the Foundation for Life Sciences (SLW), formerly the Foundation for Biological Research (BION), which is subsidized by the Netherlands Organization for Scientific Research (NWO). Cover design: Allex B. Haasdijk BI3UOTHEEK LANDBOuwuNiv;:R-rnrr CIP-data Koninklijke Bibliotheek, Den Haag Kerckhoffs, L.H.J. Physiological functions of phytochromes in tomato: astud y using photomorphogenic mutants/ L.H.J . Kerckhoffs. - [S.l. :s.n. ] ThesisWageningen .- Wit h ref. -wit h summary inDutch . ISBN90-5485-627- 0 Subject headings: photomorphogenesis / phytochrome / mutants / plant physiology / Lycopersicon esculentum ^o%?o\,z-^g Stellingen 1. De far-red light-insensitive (fri) entemporaril y redlight-insensitiv e (tri) mutanteni n tomaatzij n respectievelijk fytochroom Ae nfytochroo m Bl mutanten. 2. Inmonochromatisc hroo dlich tfungere n zowelfytochroo m Aal sB 1al shoofdrolspeler s in de anthocyaanbiosynthese in zaailingen van tomaat, terwijl micro-injectie studies alleenee nro lvoo rfytochroo m Atoekennen . Dit proefschift. Kunkelet al. (1996). PlantJ. 10:625-636. 3. Heteerde rafbreke n vanhe tontwikkelingsprogramm ai nd ehp-1 enhp-2 mutante ni n tomaatonde rcontin u verrood licht,i nvergelijkin g methe twildtype ,i see nuitermat e interessant verschijnsel, mede gezien het feit dat ook de fenotypisch gelijkende transgenetomaa twaari nhe tPHYA3 va nhave rto toverexpressi ei sgebrach tonde rdez e omstandighedenee nidentiek erespon svertoon t Dit proefschrift. 4. Ind eschaduwmijdend ereakti ehebbe nfytochroo m Bl enB 2zee rwaarschijnlij k sterk . overlappende functies. Dit proefschrift. 5. Opbrengstderving in commerciele gewassen door ongewenste schaduwmijdende reakties alsgevol gva nee nhog eplantdichthei d kunnenworde ntegengegaa ndoo rhe t viabiotechnologisch ewe gto toverexpressi ebrenge nva n fytochroom. Robsonet al (1996) .Nat. Biotechnol. 14: 995-998. 6. Dereorganisati eva nLUW-DL Otot ee nKenniscentru mWageninge ni snadeli gvoo rd e positieva nhe teerste-fas e onderwijs. 7. Deexportheffin g vand eE Ui nd ewinte r1995/199 6o ptarwe ,o mdaarme ed etarwepri js binnen de EU lager te maken dan die op de wereldmarkt, wekt de indruk dat de liberalisering van markten alleen gehanteerd wordt, zolang het gunstig is voor de consument 8. De grootste uitdaging van alle genoomprojecten is het vinden van de precieze biologischefunctie s vanall ebeschreve ngenen . 9. De beoordeling van wetenschappelijke manuscripten en onderzoeksvoorstellen door gelijken (het zgn. peer review) leidt vaak tot het beoefenen van wetenschap met voorkennis. 10. Wetenschappelijke creativiteitkom tvaa kpa sn azessen . to'&to* ,z\°\% to 11. Het is mogelijk geen toeval dathe t opheffen van deleerstoe l graslandkunde aan de LUWsamenval tme td edeplorabel etoestan d vand egrasma ti nd eAmsterda mArena . 12. Doorgeruchtmakend euitsprake n vanmgr . Muskens,zoal she ttoestaa n datarme nee n broodmoge nstelen ,i sd earmoed ei nNederlan d opd epolitiek eagend ageze t 13. Hetinbraakbeveiligingssystee mva nhe tBotanisc hCentru mi suitermat egeschik to mee n circadiaanritme t ehandhave ntijden s hetschrijve n vanee n proefschrift. 14. InMaastrich tduur td ekortst eda gee nhal f uurlange rda ndi ei nGroningen . 15. "Mijlijk tslecht s6e ndin gzeker : mensendi ebewere nda tmense ndi eva ndiere nhoude n nietva nmense nhouden ,houde nnie tva nmense ndi eva ndiere nhouden" . Koosva nZomere n(1995) ."Wa twi ld ekoe" . 16. Het veelvuldig voorkomen van de naam Kerckhoffs op lijsten van complicen en gehangenenva nd egevreesd eBokkenrijders-bend e (tweedehelf t 18eeeuw )maak td e naamKerckhoff s berucht J.H.H.Kerckhoffs ,emeritus-pastoo r(1995) . "300jaa rKerckhoffs ,ee n familie-kroniek". Stellingenbehorend ebi jhe tproefschrif t "Physiologicalfunction so fphytochrome si n tomato: a studyusin gphotomorphogeni cmutants" . Huub Kerckhoffs, Wageningen,2 0decembe r1996 . "Ofthe many intricate and beautiful control mechanisms living organisms have evolved tooptimize their survival in a variable andchanging environment, none ismore elegant than thephytochrome system ofplants" Warren L. Butler, apionee r in the study of phytochrome Voor Pien Dankwoord Ditproefschrif t ishe tresultaa t van een aantaljare n onderzoek aan fytochroom in tomaat waaraan ik met groot plezier en voldoening heb gewerkt. Hierbij zijn natuurlijk veelmense nbetrokke ngeweest .Ee n aantalva nhe nwi li kdaaro mgraa g speciaal bedanken in dit voorwoord. Alseerst e natuurlijk mijn directebegeleide r Dick Kendrick. Zijn inbreng kende letterlijk geen grenzen en ik ben hem zeer veel dank verschuldigd voor alle support, verhelderende discussies en creatieve interpretaties van de verkregen resultaten. Dicki sbehalv e inWageninge n ook in Japan werkzaam. Hij heeft mij daarom ook steeds in de gelegenheid gesteld ombepaald e lijnen van onderzoek in laboratoria indivers e buitenlanden uit tevoeren , oa. in zijn eigen lab in Japan. Daarnaast stimuleerde Dick ook het bijwonen van congressen en ikhe b dan ook vele internationale congressen kunnen bezoeken om over eigen onderzoek te rapporteren, maar vooral ook om ideeen uit te wisselen en andere mensen te ontmoeten. Ook Chris Kendrick was zeer behulpzaam met het redigeren van teksten enhandig e computertips. Jullie gastvrijheid in Bennekom was ongekend en de deur stond altijd open. Ik wiljulli e allebei bijzonder bedanken voor jullie grote inbreng, maar ook voor de gezellige tijd. Marielle Schreuder die als analist werkzaam is in onze groep, is ook zeer betrokken geweest bij dit onderzoek. Debeschreve n thresholdbox-proeven zijn doorjo u metgrot ezorgvuldighei de nprecisi euitgevoerd . Daarnaast zorgdeji j ook voor de voortgang van het onderzoek en begeleiding van onze studenten in Wageningen wanneer Dick en ik in het buitenland verbleven. Ook was jouw inbreng in de layout voor dit proefschrift groot omdatj e met veel toewijding de figuren gemaakt hebt met SigmaPlot. Marielle, enorm bedankt! Uiteraard ookmij n promotor WimVredenber g voor het mede-initieren van dit project enjou w continue interesse. Hoewelji j ons behoorlijk vrij liet, was toch ook debijdrag e in de laatste fase leerzaam en nuttig.Bedank t voor het zorgvuldig doornemen van de diverse manuscripten. Met Maarten Koornneef enAgeet h van Tuinen (vakgroep Erfelijkheidsleer) is zeer intensief samengewerkt. Jullie waren de hofleveranciers van de gebruikte mutanten enhe t was een voorrecht ome rme et e werken. Samen met Ageeth zijn ookd eeerst efysiologisch e karakteriseringen gedaanva nd ebeschreve n mutanten. We zijn beiden zo ongeveer gelijk begonnen,ji j bij Maarten en ikbi j Dick en het isee n voorbeeld van hoe interdisciplinair onderzoek elkaar aanvult en versterkt. Ook Corrie Hanhart voor het vele werk in de vermeerderings- en kruisings- programma's van de mutanten. De studenten die in het kader van een afstudeervak of stage een belangrijke bijdrage hebben geleverd: allereerst Natasja de Groot (studente Biologie, LUW) die de fysiologische karakterisering van de 'high-pigment'-achtige mutanten heeft vn uitgevoerd, waarvan de resultaten in Hoofdstuk 5 zijn opgenomen. Je hebt met veel inzet en zorg de proeven uitgevoerd. Rob van der Valk (student Tuinbouw, LUW) heeft de eerste karakterisering van diverse mutanten op vruchtniveau uitgevoerd (Hoofdstuk 6) en ondanks vele moeilijkheden in het vinden van de juiste extractie-methode, ben je er toch goed uitgekomen. Michel Sengers (stagiaire, Agrarische Hogeschool Den Bosch), die diverse mutanten heeft gevolgd metd e groeimeter (Hoofdstuk 8).Ondank s derelatie f korte tijd dieji j tot je beschikking had, heb jij zelfs nog een begin gemaakt om een automatisch watergeefsysteem te ontwikkelen. Ondanks dat ik een deel van de tijd in Japan verbleef, konden we via email goed met elkaar communiceren. Deplante n die gebruikt werden voor de groeimeter en andere proeven werden doorLee nPeters eopgekweek te nverzorgd .Aar tva nOmmere n enHen k Melissen hielpen mee met de vermeerdering van de tomaten bij ons in dekas . Jan van Kreel, Ruth vande r Laan en Ben van der Swaluw voor alle technische hand- en spandiensten waar nodig.Altij d werden verzoeken gehonoreerd en snel uitgevoerd. WillemTon k voor allehul pbi j het verder ontwikkelen van de groei­ meter, met name hetrealisere n van defotosynthetisch e belichtingen, het tunen en opnieuw uitlijnen van de 'zwevende nullen', en vele andere technische ver- beteringen die zijn ingebracht. In de meest eindeloze discussies bij jou thuis ontstonden toch ook vaak heldere ideeen en oplossingen. Rienk Bouma, Leo Herben en Chris Massop voor de ondersteuning op het gebied van de computers en software. Tot op het laatste moment heb ik kunnen rekenen op hulp waar dat nodig was. Allemaal zeer bedankt voor alle hulp en gezellige samenwerking.
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