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Quantitgaetnievtieina c psp l(eM alus domest(iBcoar kbhr.e)e)d ing: X Fruisth apter aigtesn,e tpiacr ameteesrt imatainodn breedisntgr atdeegvye lopment A thesipsr esenitnep da rtfiuallf ilomfet nhter equirements fotrh ed egroefe DoctoorfP hilosoipnhA yn imaSlc ien(cBer eedainndg Genetics) at MasseUyn iversity PalmersNtoornth NewZ ealand AlastJaoihrnC urrie 2000 Abstarct Thea iomf t htihse wsaitsso i vnestisgoamatese p eocfbt rsde ienfgoq ru antittraatiitvse ina ppleF.i rst,s tutedhxyip srl eotdh mee asuremeinntrh tieaanndc ea coofm plex quantita(tfiirvtsue h atprFeao)iu.tr a inaelry wsaissu setdom atheimcaatdlelsyc ribe appflreiu sth paei n oabjne ctmiavnen aenrpd r incciopmaplo naenanlty gsriosu ptehde Fourideers criipnttomore sai nnfugls haptert asiH.e rbiitilaeiwste rees mtaitetdo detmeirngee neltliyce ariinsthheadt prea aintgdse ,n otbyyep nev ironimneetrnaotcn tsi werees ttiemtdao d etmeirnteh set bailoiftt yht er aeixtep srsiaocnr oesnisvr onments. Friuats pecta crcaotuifnoot orev de 7r6 %o ft hpeh enotvyapriicai tnsi hoan,p f eriut conyi cif6to%ra ndfr iuts quarefnooerns ls2 y% .Th ester ahiatdms o dertaoht ei gh narro-wsenhseerb iiitltai(e0s. 709.,3a 8n d0 .3r8e spievyce)tlw,h icihn dtiectdah at inidvidsuealle cwtouilobdnee fifceinHti.hg b etwne-eesg ietnectoierclrt aio(nr> A0 .8), incdaitleodgw e noty-peen-vbiyrointnmeeronantca t nisdu ggebesretndsgi b asaetdo ne siwtoeu lbdee fifcifeonartl tesrhianapgtoe t hseir.t essAh acphea rtc owsnatsr utcot ed enabtlheet sher aepep le tsrhaatipobte ese vatleuqdau kilcayn adc ctuerilaynt hfei eld, avoiditnhngee efdoi rm gaec patuarneFd o urainaelsryi s. Seconhde,r itaabnigdle inteicteoiserc lrt aiownesre es ttiemfdao arr angea poptflr ea its froamg eneltiybc raolad-abpapsbleredde ienpgo pUlaotf7i 1fo mani liNearsr.o- wsense herilttiaywb aise smtaiteadte acohft wos itaensda corsssi tesfr iuftaoc ri d(i01t.-7y 0.2f2r)iu,ts u ga(r0s. 25-a0n.td2r 6eg)er, o whtahb (i01t9. -01.)H4.ig hehre ritabilities werees mtaitefdo frr iuts quare(n.0e30s2.s- 43fru)i,ct o nic(i.0t3-y20 .4p6o)w,d ery mildeiwcn iednc(e0 .4v0io)gu,(r 0.28-f0ur.if6tri2 m)en,s (s04 .40-.5h3a)rv,e tsitm e (06.6-.208)l,ef naig( 06.00-.8f3r)iua,ts percatt( i0o.0 7.48-f9l)o,w e(r0i.)na9 gn2 fdr iut siz(e00 .-190.1.)T raiwtista h h ihg narrow-hseernilsttieayc b oiubldes eelctfeord efifciewnittilhny d iidvusaell oen.cH tiihgg enectoierclr atbieotnw eseienst( r�A 0 6.4) inidcatleodwg enotbyyep neiv ronmietnnetr acatnitdoh nab etre ndgib asaetdo ne site woulbde e fcfiiefnotir m pvriontgh etsrea aitto st hseir.t H eisgpho sie tgievnetic coerlrtaiobnest weena tst irta1we ie trsee s taitmefdo lre ianfwgi ht floweri(n09g.5 ), harvteismte fwriiufttim hrn es(s0 .9l2e)a,fw iihnth ga rvteism(te0 .69)fur,i atac nidd ity witfhru it fi(r0m1.n.)6He ishgs n egatgievneec toierclrt aiownesr ree ptoerfdo vri gour witfhru istq uare(0n-.e8sa2sn) td r ee ghraobwitthl ewiainft(gh- 5 03.R)a.p giadni sc an bem adwei tpho sitcioverealrltyet drt asbi,u hti gnhe gatciovrerleytl raatwieitdmls al k e smiultangeaoiiunnsb otthr adiiftifsc uClatu.t miuosnt apbpel wiheedin n etrepitrng genectoircr elbaettiwotenresnad iutteso h ihgs tandearrradon rds ptohsbesl iien fluence ofe nivronmeinnettraalc tiont rbae(itdtwesep eenn daennldnci kea)dg ies ieblqriumi.u Thidr,t hter atdiionaaplpb lrede einsgta rte(gmya ssse lecwtaisoc no)ma pretdoa n opotnib aseodnr ecursreelnetc ftoigro enn ecroamlnb iinagbl iyi t(RSG-CA)T.h e excseisvlenolgyg e nerianttieoornftv hatelr atdiiobnradelie nsgt ramteeagnnyte ggliible gainwser em adHeo.w evtehrte,rd aitiborndeaielns gt ragteeengrya tgerde agtaienrps e r generatthiatonhnR e S -GCsAt rabtueatgt y g rae actoesirtnt eromfse rodgiennge tic divseirttyh,e rreebdyu ctihpneog t enftoira gll- otmne grainT.h eso lwerra toefg aiinn thReS -GCAw as tdoute hs el owreartof eg aiinnt hber enegdp ioplutaoinH.o wever, thReS G-CAs tratmeagdyeg reagtaienris n t hceu lrtsir vealattoit vhebe r enegdi poplutaiodne spsiitmeris lealecitnitoenn dsuitet obi eetsut teirtl iioosnfat hbee tween faimlgye nevtaiarcni caesw elalst hwei hti-nfmailgye nevtairci ance. Brenedgi strabtaesgoeindqe usa nttiivgteea nethiacvbsee eanp plwiietdsh u ccteos s animcarloa,pn fdo srtter berede ei,nb guotn lrye ceinnta lpypb lree negdI.it is clear that ftuurber eedpirnogg rawmomueslb dee nffirto tmh aep plicoafqt uiaotnnat tigievenet ic theory. if Acknoweldgemnets Thitseh sicsou lndo t havmea dbpeeo esns iwbilteht ohupeta tieansscitesa,n acned encouragfremom ewani tdr ea nogfpe e ople. Ia mg ratteofPu rlo Dfo.r iGaanr r(iMcaks sey Univcehrisseiuftp ye)r,vf iomsryo r, takionngy eta nothsetru deinnh ti she avy worIk hlaovaaedp .p rechiiatse d encoguermaeanntdg uidatnhcreo tuhgtehh eoorfby r eiendga,n dh icsl re,ca oncise cirticfioesram cp hrj oect. Ia mg reaitneldbyt teoDd r D.o minuieNq oit(oHno rtReswehaowr acsfhi )rm sybt o ss, themny s upervwirsiotrti,un tgao nrda lwaay fsre indD.o mniiuqed evelotpheed quantigteanteiptvrieoc gsr aamnmsdee tt husepc hroslhatioep n abmlete o p ursfuluel timset udTyh.r ougthhoeul ta sftwe yearssh e bheaesan cosntasnotu rocfe encouraggeumiednaatnn,cmd eo tvitai.o naIl saop prectihmaeot rere e ceenftf oorft s DrN.n daozOirea gu(zHioer tRcehs)we,ha osr upplied avdavlioucnate bh wlere i gt in ansdrt uctouftr hete h easswi es lalso fefringu lsh ueglgpefsftorei aocnchsh apter. Ia mt haunlfk ofDrr T.o nSyhe lboeus'rn( ForReesste airnclvhua)ba liepn uittn o every aspeocftt hs ti ehsiHsei. nr todumceet dot hfeo rtersebter enegdt iheotrhyat th is thesis isfo nudeodn I.h ave baelnseoff iortmte hdie n poufot t hberre edaetFr osr eRsetsr ecah (DrM.i kCea rsDorn.R, o wlaBnurdod nD,r L. uis DGre.Pa a,u Jlef efrsoSnio,mn WeaveDrr,.K eitJha yawriacmkaa nd DSrue. C arsoTnhe)i.ir n puhta sb een eduicoantmaolt,i vaatngidrn ega atplpyir eacted. Thisteh siwosu lndoh ta vbee epno sbslwiei thaocuctet sofsn i naclis aupoprftca,ii iltes andt haep pblree negdp irograomfemfree bdyT heH orticualntdFu oroaRdle saerch Insutti(etH ortRcehs)Ne,ea wrZ ealaInnpd a.r tiIct uhlaanrk P aDurGl l uciDnra., DominiNqouieta onnd NDnra.d oOzriaeg u(zmiaen ageamnedMn tad)e leHionf stee anJda niFcrea s(etre chansiscaianslct e). Freaec cetsoAs r thGuimrlo u'rsA SREML proagnrdha me luplsf uggeswteiroen s appraetcefido r etshteti imooafng eneptraiacm e.tT ehrems aps wgeernete erwdai tthh e iii map-makpirnogg raavma ilaobnl teh eC harlSetsu rUtn iverwseibts yi te ht:t//pilef.c.seu.duauc/g-ibin/gis/Map. Througmhyot uitma etM assIeh yav eb eelnu ctkohy a vgeo ofdre indasn cdo lglueeas thhaatv per oviqdueeidsno tsa,ns werfsee,d baancddki verTshiaonynko! eu s pieacltloy SatisKhu maLru,i Asp izoalL,ai sWaa tsoNni,c< oiVlsi llloabaonsd t hree sotft he posatdguraitne sr 2o.o0m2 . AlhtougaPh h Ds tudheasnl tti lter ets ipmaIet ,h amnekm beorfSs tA lbaPnrse sbyterian Churfcohsr t crheitmnygh orizboenyso tnhdhe e raen dn o.wT hey hparvoev ided balainnmc yel eia fnhde lpmeed ftocou osn wihrsae tla yil mportant. Lasyt lwIo udl likteog ivaes pecaicakln owledtgome ymf emanit,l e yscpieamlyl y wifeD,e borfaorlh o, vaens du prtp tohroughout. Twoa re btehtaotnne ebr,e catuhseheya vaeg oorde tfuorrnt ehiwro kr: Ifo nfeal ldso wnh,if sri encda hne plh iump . Buptiy tt hmea n wfahlola sn d nhoao sn teo h eplh imu p! Eec 94-:1 0 iv Contents ABSTRACT 1 ................................................................................................................................................ ACKNOWLEDGEMEN..T...S... ..........................................................................................................I.. II CONTENT..S... ..........................................................................................................................................V LISOTF T ABLEASN DF IGURES VI ....................................................................................................... CHAPTE1R: I NTRODUCTION SUMMARYO FC HAPTERS 2 .................................................................................................................... CHAPTE2R .RE VIEWO FL ITERATUARpEP:L EB REEDINSGT RATEGAYN DO BJECTIVES . 2 .......... ................... CHAPTE3R .Ap PLES HAPTER AITS ..... . .. 3 ............. .... .............. ............ . ............................ ............................. CHAPTER E4S.T IMATIGNEGN ETIPCA RAMETERS .. . . 4 ........... ............. ............. ....... .................. ................... CHAPTE5R. Ap PLBER EEDINGS TRATEGY . 5 ..................... ...................................... ..................................... REFERENCES 6 ........................................................................................................................................... CHAPTE2R: L ITERATURREEV IEWO FA PPLEB REEIDNG INTRODUCTION 8 ................................................................................................................................. ..... GENETIOCR IGIONF T HEA PPLE 8 ...................................................................................................... TAXONOMYO FA PPLE .. . . . 8 ............................... .. ........................ ..... .... ....................... ................................. SPECIEWSI THITNH EM ALUSG ENUS. .. . . . . . . 9 . . .. ...... ........ ...... ..... .................. ... ......................... ..................... GEOGRAPHIDCI STRIBUTOIFOM N. X DOMESTlC.A.... ..........................................................1...1 BREEDINSGT RATEGIFEOSR A PPLE 12 ............................................................................................. INITIBARLE EDINSGT RATEGIES . 12 ................. ......................... ... ................................................................. CONTROLLEPDO LLINATION .. . . 14 ............................... . ........ .................................................... .................... CURRENT BREESDTIRNAGT EGIES. . . .. . .1 4 ..... ............. ........... ............ ........... ........................ ........................ Recurrmeanstss e lecti.o.n . ... . . . 15 ........ ...... .... . ... ......................... .............. ........................ ..................... Recurrseenlte cftoiGroC nA ( RS-GC..A) . . . . 15 .. ......................... ..... ............................ ................. ......... Inbreeding . . 17 ............... .... ................................................................................. ................................... Modifbiaecdk cross . .. . 18 .............................. ................................. ...... . ................................... .............. Markeasrs issteelde cti.on ... . 19 ..... .......... ................... ... . ............................................... ........................ Genter ansform.a.tion . . .. . . . .2 1 .... ................... ..... ............. .. ..... ....... ........... ................ . ............................ Mutation breedin.g . . 21 .................... .............. ...................... ...... ............................................... ............. BREEDINOGB JECTIVAENSD S ELECTICORNI TERFIOAR A PPLE 23 ....................................... WHATA REB REEDINGO BJECTIVAENSDS ELECTIOCNR ITERI.A.? . . ... 23 ... ............ ......... ...... . ........................ MULTIPLE TSREALIETC TIIONNA PPLIEM PROVEMENT . . 23 ....................... ....................... ....................... ..... DEVELOPMEONFTB REEDINOGB JECTIVAENSDS ELECTICORNI TERIINAA PPLE 24 .............. ........................ CURRENDTE SSERATP PLBER EEDINGO BJECTIVES .. . .25 .................................. . ............ .............................. .. Fruqiuta l.ity . . ... . . . 25 ........................... .... ....... .. ... ........ ............. .................................................... ......... Producctoisso tn . . 30 ....................... ............................................ ........................... ................................ Productivi.ty. . . . .. . 34 ........... . .................. ...... ..... ..................... .. ...................... .......... ............................... SELECTIIONND EX . . 37 ...... ................ ..................................... ........... ......................... .................................. SUMMARy 38 ............................................................................................................................................... v REFERENC.E...S.... .................................................................................................................................3 8 CHAPTER 3: QUANTITATIVEE VALUATIONO F APPLE (MALUS XDOMESTICAB ORKH.)F RUITS HAPE BY PRINCAILP COMPONENT ANALYSIOSF F OURIEDRE SCRIPTORS ABSTRAC...T... ........................................................................................................................................5 6 ABBREVITAI ON..S... ..............................................................................................................................5 7 INTRODUCTI...O...N... ...........................................................................................................................5 7 MATERIALASN DM ETHOD.S... ......................................................................................................... 58 VISUALISATOIFOP NR INCIPCAOLM PONEN(TP CS)H APE TR.A.I..T.S... ..........................................5..8.. . . . . . . . GENETIACN ALyS.I.S. ...................................................................................6...0... ...... ............................... SHAPE CHART D..E.S.I.G...N.. .................................................................................................6..1 .. . .. .. ... . . . . . . RESULTS 62 .................................................................................................................................................. VISUALISATOIFOP NC S HAPET RAIT.S. ............................................................................6..2... ....... . . . . . . . . . . . GENETIC ANAL.y.S.I.S. .....................................................................................................6...2 .. ... . . . .. . . . . . . . . . . INTERPRETATOIFPO NC T RAIT.S. .........................................................6...3... ....... ..................................... SHAPEC HART. ..........................................................................................................................6. 4. .... . . . . . . . . DISCUSSI...O....N... ..................................................................................................................................6 4 ACKNOWLEDGEMENTS . . . .67 ................................................................................................................. REFERENCES . .. 67 ...................................................................................................................................... CHAPTER 4: ESTIMATESO F HERITAIBLITIESA ND GENETIC CORRELATIOFNOSR A PPLE( MX.D OMESTIBCAO RKH.T)R AITS ABSTRAC.T. .........................................................8..0.. ............................................................................... INTRODUCTION. . .. . 80 ............................................................................................................................... MAT ERIALASN DM ETHODS . . . . . 84 ........................................................................................................ EXPERIMENTADLE SIG.N... .........................................................................................................8..4. .... . . . .. . MODELE QUATIO.N. ...............................................................................................8.4. .............. . .. .. . . .. . . . . .. . . HERITABIL.I.T..Y.. ..............................................................................................................8.6. ....... . . . .. . . . . . . . GENETICCO RRELATI..O.N. .....................................................................................,.... ... .......8.8. ....... . . . . . . . .. RESULTS . . 88 ................................................................................................................................................ ESTIMATIVNAGR IANCCEO MPONEN.T.S. .................................................................................8..8.. ..... . . . . . . . HERITABILI..T..y.. ........................................................................................................8 ..9.. ...... .. . . . .. . . . . . . . . . . GENETICCO RRELATIBOENT WEESNI TE.S. ........................................................................8.9. ................. . . . . GENETICCO RRELATIBOENT WEETNR AIT.S. ..............................................................................8...9 . .... . . . . . . DISCUSSION . . 90 .......................................................................................................................................... REFERENC.E.S. ................................................................................9...5.. ................................................. Vl CHAPTER5 :C OMPARISOONF T WO BREEDINSGT RATEGIIENS APPL(EM . XDOMESTICBAO RKH.) ABSTRACT 104 ............................................................................................................................................ INTRODUCTION 104 .................................................................................................................................. TRADITIONAAPLP LBER EEDINSGT RATEGY I07 ........................................................................... TRADITIONBARLE EDING POPULSATTRIUOCNT URE 107 ............. .......................... ....................................... TRADITIONMAALT INDGE SIGN 108 ........................................ ..................................................................... TRADITIONSAELL ECTION 108 .............................................. ..................................... .................................. RS-GCSAT RATEGY 109 ............................................................................................................................ RS-GCBAR EEDINPGO PULATION 110 ......................................................................................................... RS-GCbA reedpionpgu lasttirounc ture 110 .......................... ................................................................ RS-GCbA reedpionpgu lation mating design 111 ........... ....................................................................... RS-GCBA reedpionpgu lasteiloenc tion 113 ................... ................................... .................................... RS-GCCAU LTIVAPRR ODUCTIPOONP ULATION 114 .............. ............................ .......................................... RS-GCAc ultipvraord ucptoipounl asttirounc ture 115 .......................................................................... RS-GCAc ultipvraord ucptoipounl amtaitoindn egs ign 115 ...... ........................ .................................... RS-GCcA ultipvraord ucptoipounl asteiloenc tion 115 .......................................................................... SIMULATION 116 ........................................................................................................................................ SIMULATIMOENT HOD 116 ........................... ............................... ................................................................. Traditbiroeneadlsi tnrga tmeogdye l 117 ................................... .................................. .......................... RS-GCbA reedsitnrga tneTgoyd el 118 .................................... ............................................................... SIMULATIROENS ULTS 118 ......................... ................................. ..................... ............. ............................... SIMULATIODN! SCUSSION 119 .................... ........................... ....................................................................... CONCLUSIONS . 120 ....................................................................................... ............. ................................ REFERENCES 120 ...................................................................... ................................................................. CHAPTER6 :G ENERADLI SCUSSION REFERENCES 136 ....................................................................................................................................... vii Lisotf f iguraensd t ables Chapte2r: L itetruaroef a pplber eeding Figu1r.De ia graomft hpeo sseei vbolluotfti hoReno sacseuabef laeimsbi asoend moroplhogiccyatlo,l oagncidhc eamli dcaat(laC hiacl1el9, 7) 4 50 . ........................... Figu2r.We o rld omfta hpoe ir gionfas p pslpee cies. . . 51 ............ ........ ........................ ....... Figeu3 r.Wo rld omfta hpoe ir gionfs aspppelcei inoevvlse idn t haen ceosft rtyh e domesatplipec .. . . 52 .................... ....................... ........................ ......................... .... .... Figu4r.Me ap o ft hdei striabnudt idoonm esotfai pcpa(ltMeiX. o d no mesti.ca5)3 ...... .... Tab1le.O r giiann bdo tandiecsaclr ipatpipsolnpe ee ocsifi vn olvientdh d eo mestication ofa ppl.e . . . . . .54 .. ............ ................ ..... ............ ...................................... ............... ...... .. Chapter3 :Q uantittaiveev aluatioofna ppl(eM aluxs d omestica Borkhf.r)u it shbaypp rei ncipcaolmp onenatn aylsiso fF ouirer descirptors Tab1le.P r optoiroofnt hteo tpahle notvyapriicaa nccceo ufnotre tdbh fyei r6Ps Ct.s 7 1 Tab2l.eV isulaitsiaoofnt hferi sstiP xC s hatprea (imtimsnu imt o miamuxmv ael)u. Appslhea paerdser awwni tthhs et eemn adtt h teo apn tdh ceay lxa tt hbeo tt7o2m . Tab3l.eH erbiiityltae sttiemsa s(atannddae rrdr ofroPsrC) a pple tsrhaaaitptt ehs e HavelNooctrkha nNde lssoint es . . 73 ............................ ......................................... .... Tab4l.eG enectoiercalrt io(rnbg e)t wseietnae nssd t arndde rarfoorar p ple tsrhaaiptes 74 ............................................................................ .................................................... Tab5l.eR egersscioofenif ciebnettsw eenP Cas phppaelt er aaintcdsla leirp measurements .. .. 75 .......................... ....... ... .................... ........................................ ...... Tab6l.Ae p pslhea cphaer bta soenda speccotn,i acnisdtq yu arePnCt ersta.ssAi pple shapaerdser awwni tthhs et eemn adtt he atnotdph c eay lxa tt hbeo ttom 76 ........... Figeu1 r.Ap pclreo ss-ssehcontwgiti ohcnea lelrmi epasuernetBms =. s teGm =, c alyx, BG= stem-caaxi,lsC yHx= lnegt(hL D)E, = width El(= Wm)a,x imwuimtd h dsitan(cMeW D)A,C= stecma vwiitdytF Hh= , c aylxb aisnw id.t..h. .......79.. ......... Chapter4 :E stiamteso fh eritlaiybt iandg eneitcc orreltaiofno r appl(eM aluxs d omesticBao rk.h)t riats Tab1le.N a mea ndde scrifpoetrai coahnp ptlre.a .i.t. ................9.9. ................................. Tab2l.Be asisct atiforsa tpipctlsre a aitHt asv elNoocrk(t ihst 1e,)N eslo(ns ei2 t)a nd combisnietodef ts hb er eiendpgo pulation. . 100 ................. ................................... ..... Tab3l.eU nirvaiatnera ro-wsehnesrebi ityla( itstandearrrdio nbr r ackfeoatrsp )pt lrea its esttiemsdae rpaaltyfeo r HaveloacnNkde loNsnoa,rnt adhc robsosts hi .t.e.s1. 0.1. . viii
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