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Synthesis and Fertility of Xbrassicoraphanus and Ways Of i C?2o Q\7- C Synthesis andfertilit y of xBrassicoraphanus andway s of transferring Raphanus characters to Brassica O.Dolst i NN0B201.912 O. Dolstra Synthesis andfertilit y ofxBrassicoraphanu s andway so ftransferrin g Raphanus characters toBrassic a Proefschrift terverkrijgin gva nd egraa dva n doctori nd elandbouwwetenschappen , op gezagva nd erecto rmagnificus , dr.C.C .Oosterlee , hoogleraar ind eveeteeltwetenschap , inhe topenbaa rt everdedige n opvrijda g2 9oktobe r198 2 desnamiddag s tevie ruu ri nd eaul a vand eLandbouwhogeschoo l teWageningen . Centre for Agricultural Publishing and Documentation Wageningen - 1982 Abstract Dolstra,0. ,1982 .Synthesi san d fertility ofxBrassicoraphanu s andway s oftransfer ­ ringRaphanu s characters toBrassica .Agric .Res .Rep . (Versl.landbouwk .Onderz. ) 917,ISB N90-220-0805-3 ,(viii )+ 9 0p. ,3 8tables ,1 4figs ,11 4refs ,Eng .an dDutc h summaries. Also:Doctora lthesis ,Wageningen . About25 0intergeneri chybrid swit hth egenom e constitutionAR ,AAR Ro rAR Rwer eob ­ tained fromove r 1500 0crosse sbetwee nBrassic a campestris (AAo rAAAA ,femal eparent ) andRaphanu s sativus (RRo rRRRR) .Th epoo rcrossabilit ywa s shownt ob ea consequenc e ofvariou sbreedin gbarriers .Th ediploi d andtetraploi dhybrid swer e studied inform , fertility,chromosom eassociatio na tmeiosi san d crossabilitybot hamon gth ehybrid s and incrosse swit hth eparenta l speciesan dBrassic a napus•XBrassicoraphanu s (AARR) wasbackcrosse d twicewit hB .campestri s (AA),an dwa sals ocrosse d andbackcrosse d withB .napu s (AACC).For man d chromosomeassociatio na tmeiosi s inAA Ran dAAC Rhy ­ bridswer estudied . Twopopulation so fXBrassicoraphanu s (AARR)wer epropagate d and studied inmor ede ­ tail,partl y forpossibl esuitabilit ya sa ne wcrop .Mos tplant swer ehighl ysteril e becauseo fbreedin gbarriers ,whic hwer esimila rt othos ei nth eorigina l intergeneric crosses.Th eincreas ei nfertilit yan dchromosoma l stabilitywer e studied inth efirs t threegeneration s ofth epopulations . Freedescriptors :Brassic a campestris,Raphanu s sativus,incongruity ,breedin gbar ­ riers,embry oculture ,intergeneri chybrids ,XBrassicoraphanus ,fertility ,matin g system,introgression ,Brassic anapus ,allosyndesis ,forag ecrop . Thisthesi swil l alsob epublishe d asAgricultura l Research Reports917 . © Centre forAgricultura l Publishing andDocumentation ,Wageningen ,1982 . Nopar t ofthi sboo kma yb e reproduced orpublishe d in any form,b yprint , photoprint,microfil m oran y othermean swithou twritte npermissio n from thepublishers . Stellingen 1.Selecti e in Brassica campestris en Raphanus sativus opee ngoed ekruis - baarheid tussen deze soorten,i nhe tbijzonde r volgens demethod eva n de reciproke récurrente selectie,za lnie t alleen leiden tot eenverbeterin g vand ekruisbaarheid , maar ook totee nverhoogd e fertiliteitva nd e allo- tetraploïde geslachtshybriden. Ditproefschrift . 2.Mannelij k sterielehomozygot e lijnenva nkoolzaa d enbladkoo l zijnver ­ krijgbaar uitheterozygoo tmannelij k steriel uitgangsmateriaal doorher ­ haalde inductieva n diploïdeparthénogenès e (matromorfie)vi abestuivin ­ genme t xBrassicoraphanus. Ditproefschrift . 3.Polyploïdisati e doorvalentiekruisinge n (4xx 2 xkruisingen )i soo kbi j generatiefvermeerderd e gewassent everkieze nbove nmitotisch everdubbe ­ lingva nhe tchromosoomaantal . 4.He t iswenselijk , datsynthetisch e amphi(di)ploïde vormen diebruik ­ baar kunnenzij nbi j deveredelin g van cultuurgewassen wordenbewaar d in genenbanken, ookwannee rhe tnu thierva nnie tdirec tduidelij kis . 5.Te nonrecht ebeschouw tRöbbele n allopolyploïdie alsee n doodlopende weg ind eevoluti eva nplanten . G.Röbbelen ,1979 .Proc .Conf .Broadenin gGenet .Bas eCrops .Wageningen ,1978 . Pudoc,Wageningen ,p .249-255 . 6.D everedelin g vanmai s zou zeergebaa tzij nbi j openbaarheid van de formulesva nhybrid e rassen enbi jee nregelin g diehe tmogelij k maakt datd eouderlijne nva nhybrid e rassenbeschikbaa r komenvoo rwetenschap ­ pelijk onderzoek en kruisingsdoeleinden. 7.He t isnie twaarschijnlij k dati nNederlan d de teeltva nmai svoo r 'corn-cob-mix'ee ngrot evluch t zalnemen . 8. De plantenveredeling zou verrijkt worden met een aantrekkelijke methode voor het verbeteren van waardevolle genotypen, indien somatische recombi- natie in een relatief hoge frequentie geïnduceerd zou kunnen worden in cultures van protoplasten en cellen. 9. De 'recombinant-DNA'-technieken vormen weliswaar een belangwekkende nieuwe technologische ontwikkeling, maar de betekenis ervan voor de plantenveredeling wordt erg overtrokken. R. Keuning et al., 1980. Chemisch Magazine. Bijvoegsel Chemisch Weekblad 10: 156-161. J. Ingle, 1982. Span 25: 50-53. Proefschrift van 0. Dolstra characters' £££* °f ^-^^or^^s and ways of transferring Ra^hanus Wageningen,2 9oktobe r198 2 Woord vooraf Hetbeschreve n onderzoekwer dverrich t aan de Stichtingvoo rPlanten ­ veredeling (SVP)t eWageningen , inhe tkade rva nhe tprojec t 'Cytogene- tischonderzoe k aansoort -e ngeslachtsbastaarde nva nlandbouwcruciferen' . Hetbestuu r end edirecti eva n de SVPhebbe nmi j in staatgestel d dit onderzoek teverrichte n end e resultaten tebewerke n totee nproef ­ schrift. Ikbe nhe n zeer erkentelijk voor hetvertrouwe n datz e daarbij inmi jgestel dhebben . Ditproefschrif t isto tstan dgekome nn a intensief overlegme tprof . dr. ir.J.G.Th . Hermsen enprof .dr . ir.J . Sybenga.Hu n opbouwende com­ mentaren hebben eenduidelijk e invloed gehad op de eindversie van dit proefschrift. Ikbe nhu nhiervoo rvee l dank verschuldigd. Eenpromotie-onderzoe k isnooi tee n eenmansactie.Vee l SVP-ers hebben hieraan eenzee rgewaardeerd e bijdrage geleverd. Enkelenwi l ikhie r met name noemen. Ir.H .Toxopeu s heeftee ngrot e inbreng gehad bijhe t formu­ lerenva nbovengenoem d project enveelvuldi g alsvraagbaa k gefungeerd. Zijnenthousiasm evoo r de landbouwcruciferen werktebijzonde r stimulerend. Verderwer dvee l steunverkrege nva n ing.G.J . Speckmann en ind e laatste jarenvoora lva n dr.ir .W . Lange.D e heerJ.H.M .Zuidgees theef t ind e eerste jarenva nhe tonderzoe k opbekwam e en actievewijz e assistentie verleend. Ind e slotfaseva nhe tonderzoe k hebbend ehere n J.H. Lubberts enH .Dijkstr a mettoewijdin g informatie verzameld over de fertiliteit van xBrassicoraphanus. Kortetij d zijnvoort s de STOVA-stagiaires Marianne van derBlom ,Aleid ava n deKraat s enGerar d Bremer actiefbi jhe tonder ­ zoekbetrokke ngeweest . Hetvel e typewerk iso p de SVPuitgevoerd , waarbij inhe tbijzonde r mw.A .Meereboe r zichverdienstelij k heeftgemaakt .Vee l foto's in hetproefschrif tzij ngemaak tdoo r fotografenva nhe tBG D teWageningen . De grafiekenwerde ngeteken d door dehee rG . Post (SVP).D e correctie van deEngels e tekste nd euitgav eva nhe tproefschrif twerde n respectieve­ lijkverzorg d door dehee r J.C. Rigg B.Sc. en ing.J . Casteleinva n het PUDOCt eWageningen . Curriculum vitae De auteurwer d op2 6 januari 1946 te Steenwijkerwold geboren. In196 5 behaalde hij aanhe tJ.H . TrompMeesterslyceu m te Steenwijk het diploma HBS-B.N ahe tvervulle nva nzij nmilitair e dienstplicht,bego n hij in 1967ee n studie aand eLandbouwhogeschoo l teWageningen ,waa rhi j inja ­ nuari 1975he tdoctoraaldiplom a behaalde ind eplantenveredelin g met als keuzevakken erfelijkheidsleer (verzwaard)e n algemene planteziektenkunde. Sinds 1 september 1974 ishi j indiens tva nd e Stichtingvoo r Plantenver­ edeling teWageningen ,waa r sindsdien hetbeschreve n promotie-onderzoek werdverricht . Ind eperiod e 1974-1977wa shi j tevensverantwoordelij k voor het cytogenetisch onderzoek aangranen . Sinds eind 1977 is de auteur belastme the tveredelingsonderzoe k aansnijmais . Contents 1 General introduction 1 1.1 Rationale 1 1.2 Speciesan dallopolyploi dhybrid s 1 1.3 Scopeo fth eresearc hprogramm e 2 2 Crosses between Brassica campestris and Raphanus sativus 5 2.1 Introduction 5 2.2 Materialan dmethod s 6 2.3 Results 9 2.3.1 Settingo fsiliqua ean dseed s 9 2.3.2 Seedsiz e 9 2.3.3 Productiono fhybrid san dmatromorph s 10 2.3.4 Useo fgeneti cmale-sterilit y 13 2.3.5 Cultureo fembryo s 14 2.3.6 Genomeconstitutio nan dnumbe ro fchromosome si n theprimar yhybrid s 15 2.4 Discussion 17 3 Nature of breeding barriers 19 3.1 Introduction 19 3.2 Materialan dmethod s 19 3.3 Results 20 3.4 Discussion 22 4 Primary allodiploid ARhybrids and their progeny 24 4.1 Introduction 24 4.2 Materialan dmethod s 25 4.3 Results 26 4.3.1 Formo fA Rhybrid s 26 4.3.2 Chromosomeassociatio na tmeiosi s 28 4.3.3 Malean dsee dfertilit y 30 4.3.4 Crossabilityo fA Rhybrid swit hvariou sspecie s 31 4.3.5 Somatican dmeioti cdoublin go fchromosom enumbe r 33 4.3.6 Progenyo fcrosse sbetwee nA Rhybrid s 33 4.4 Discussion 34 5 xBrassicoraphanus: form, fertility and crossability with various species 37 5.1 Introduction 37 5.2 Material andmethod s 38 5.3 Results 38 5.3.1 Form and fertilityo fprimar y hybrids 38 5.3.2 Crossabilitywit h Brassica campestris, Raphanus sativus and B. napus 40 5.4 Discussion 42 6 xBrassicoraphanus: sterility factors and mating system 44 6.1 Introduction 44 6.2 Material andmethod s 45 6.3 Results 45 6.3.1 Meiosis 45 6.3.2 Barriersi nth eprogami c phase 49 6.3.3 Barriers after fertilization 52 6.3.4 Mating system 54 6.4 Discussion 55 7 Improvementin fertility of xBrassicoraphanus 57 7.1 Introduction 57 7.2 Material andmethod s 57 7.3 Results 59 7.3.1 Chromosome numbers 59 7.3.2 Changesi nfou r fertility characteristics 60 7.3.3 Effectso fchromosom e numbero nfertilit y 64 7.4 Discussion 64 8 AAR hybrids and introduction of alien chromosomes into
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