GENETIC VARIABILITY FOR GIRTH GROWTH AND RUBBER YIELD IN Hevea brasiliensis

46 Gonçalvs t al. GENETIC VARIABILITY FOR GIRTH GROWTH AND RUBBER YIELD IN Hva brasilinsis Paulo d Souza Gonçalvs 1 ; Marlo d Almida Silva ; Ligia Rgina Lima Gouvêa 1 ; Erivaldo José Saloppi Junior 3 1 Apta/IAC - Programa Sringuira, Av. Barão d Itapura, 1481, C.P. 8-13001-970 - Campinas, SP - Brasil. Apta/DDD - Pólo Rgional Cntro Ost, C.P. - 1701-970 - Jaú, SP - Brasil. 3 Apta/DDD - Pólo Rgional Norost Paulista, C.P. 9-15500-000, Votuporanga, SP - Brasil. Corrsponding author <paulog@ia.sp.gov.br> ABSTRACT: Basi knowldg of gnti haratristis of populations is nssary to ondut fftiv brding and sltion. Th objtiv of this papr is dsribing th gnti variation of rubbr yild and th orrlation with othr traits, and stimating th gnti paramtrs foirth growth and total numbr of latx vssls. Sixty svn lons of Hva brasilinsis (Willd. x Adr. d Juss.) Mull.-Arg. wr tstd at fiv sits during 10 yars. Charatrs girth growth at panl opning and rubbr yild, showd broad sns hritability on plot man lvl, from 0.3 to 0. and 0.59 to 0.9, rsptivly. Prditd gnti gains qual to 0.73 m and 0.79 g inras rsptivly on girth and yild in th opning panl and matur phass sms ralisti, vn with modrat sltion intnsitis. Gnti orrlations with rubbr yild, bark thiknss and total numbr of latx vssls wr vry larg, and almost no gnotyp-nvironmnt intration was prsnt foirth growth. High gnotyp-nvironmnt intration was prsnt for rubbr yild with gnti and phnotypi orrlations aross th sits, ranging from 0.64 to 0.9 (gnti) and 0.63 to 0.89 (phnotypi). Total numbr of latx vssls rings had a high hritability, ranging from 0.0% to 64.0% in th sits E and B, rsptivly. Ky words: rubbr tr, hritability, gnti orrlations, gnotyp-nvironmnt intration VARIABILIDADE GENÉTICA PARA CRESCIMENTO DO CAULE E PRODUÇÃO DE BORRACHA EM Hva brasilinsis RESUMO: O onhimnto básio das aratrístias prsnts nas populaçõs d plantas é ssnial para a ondução dos trabalhos d slção mlhoramnto. Nst trabalho, objtivou-s dsrvr a variação gnétia da produção d borraha orrlaioná-las om outras aratrístias, assim omo stimar os parâmtros gnétios para prímtro do aul o númro d anéis d vasos latiífros. Um total d 67 lons d sringuira [Hva brasilinsis (Willd. x Adr. Juss.) Mull.-Arg.] foram avaliados m ino loais durant 10 anos. Os aratrs prímtro do aul na abrtura do painl produção d borraha mostraram hrdabilidads no sntido amplo ao nívl d média d parla ntr 0,3 a 0, 0,59 a 0,9, rsptivamnt. Ganhos gnétios prditos d 0,73 m 0,79g d inrmnto d prímtro na fas juvnil parm viávis msmo om uma intnsidad d slção modrada. Corrlaçõs gnétias ntr produção d borraha, spssura d asa númro total d vasos latiífros foram muito altas, pratiamnt não houv intração gnótipo-ambint para prímtro do aul. Alta intração gnótipo-ambint foi dttada para produção d borraha, om orrlaçõs gnétias fnotípias por loais variando ntr 0,64 0,9 (gnétias) 0,63 0,89 (fnotípias). O arátr total d anéis d vasos latiífros aprsntou alta hrdabilidad, variando d 0% até 63% nos loais E B, rsptivamnt. Palavras-hav: sringuira, hrdabilidad, orrlação gnétia, intração gnótipo-ambint INTRODUCTION To improv th produtivity of rubblantations, Hva brding programs xploit gntially variabl populations to obtain suprior trs. A basi knowldg of th gnti haratristis of plant populations is nssary to ondut fftiv brding and sltion. Quantitativ information is rquird about th siz of gnti varians, th typ of gn ation, and th hritability and gnti orrlations for onomially important traits. This nabls th outom of sltion to b prditd, partiularly gnti gains. It also hlps dtrmining diffiultis in sltion and th stratgis to ovrom suh problms. In a broadr ontxt, it broadns knowldg of th gntis and brding bhavior of th spis involvd. Th Instituto Agronômio (IAC) onduts a Brazilian Hva brding program, with mphasis on Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

Girth and rubbr yild in rubbr trs 47 improvmnt and sltion of vigour and quality traits, suh as latx vssls rings. Th objtiv of this papr is dsribing th gnti variation of rubbr yild, and its orrlation with othr traits. Gnti paramtrs foirth and total numbr of latx vssls of 67 lons, 13 yars aftr stablishmnt in th fild, wr also studid. MATERIAL AND METHODS Plant matrial and xprimntal loations Thirty on Brazilian, 10 Indonsian, and 6 Malaysian Hva brasilinsis gnotyps (lons) wr usd in this study. Th Brazilian lons Fx, IAC, and IAN wr dvlopd by th Fordland brding program, Instituto Agronômio (IAC), and formr Instituto Agronômio do Nort (IAN), now Embrapa Oidntal. Th Indonsian lons omprisd GT, PR, and AVROS, from Gondang Tapn, Profstation voor Rubbr, and Algmn Vrning Rubbr Plantrs Oostkust Sumatra xprimntal stations, rsptivly. Th Malaysian ultivars onsistd of RRIM and PB, from Rubbr Rsarh Institut of Malaysia, and Prang Bsarivat rubblantation rsptivly. Among th 67 lons, PB 35, GT 1, RRIM 600 and IAN 873 ar rommndd prodution lons, whras RRIM 701, PR 55, PR 61, IAN 873, PB 16, and RO 38, ar usd as parnts in loal rubbr brding programs. Th rubbr lons wr graftd ovr stablishd GT 1 rootstoks at th nursry. On yar-old rootstoks sdlings raisd in nursris wr usd to budgraft lonal matrials. Th sussful budgrafts wr uprootd and plantd in polythyln bags. Aftr th first flush of lavs dvlopd, plants wr stablishd in th fild. Th lons wr grown for tn yars in fiv ontrasting tst nvironmnts (A, B, C, D and E) in th platau rgion of São Paulo Stat, ologial onditions summarizd as: Jaú (A): º17 S, 48º34 W; altitud 580 m; man annual tmpratur 1.6ºC; man annual rainfall 1,344 mm; Kandiudox soil, with good nutrint status and physial strutur. Mooa (B): 1º18 S, 47º01 W; altitud 5 m; man annual tmpratur 4ºC; man annual rainfall 1,500 mm; Eutrustok soil, with good nutrint status and physial strutur. Ribirão Prto (C): º11 S, 47º48 W; altitud 467 m; man annual tmpratur 9ºC; man annual rainfall 1,530 mm; Kandiudox soil, with good nutint status but poohysial strutur. Votuporanga (D): 0º5 S, 49º50 W; altitud 450 m; man tmpratur during growing sason 3ºC; man annual rainfall 1,480 mm; Paludalf soil, with avrag nutrint status and poohysial strutur. Matão (E): 1º18 S, 48º40 W; altitud 551 m; man annual tmpratur 5ºC; man annual rainfall 1,480 mm; Paludox soil typ, with avrag nutrint status and vry good physial strutur. Ths loations rprsnt th most important ontinntal limat, non-traditional rubbrodution aras in Brazil. Exprimntal dsign at ah tst loation was randomizd omplt bloks (n = 3), with tn plants pr on-row plot in all loations. Trs wr spad 8.00 m btwn and.50 m within rows (500 trs ha -1 ). Masurmnts At th nd of th svnth yar, girth, bark thiknss and total latx vssl rings at panl opning for tapping and rubbr yild of ah tr wr masurd. Girth masurmnts wr rordd at 1.0 m from th highst point of th bud union. For annual latx prodution, attmpts wr mad to rord fiv annual yilds aftanl opning. Th latx xtratd from th tapping panl followd a half-spiral, four-daily tapping systm (svn tappings pr month), 11 months pr yar. Aftr tapping, latx was olltd in plasti ups providd for ah rording tr. On th latx flow was stoppd, rubbr was oagulatd in th up by adding % ati aid solution and stirring. Th oagulatd rubbr was prssd into a ylindr, hang-drid for 30 days, and thir wighd for alulation of th dry rubbr ontnt. Statistial analyss within sits Th following linar modl was usd to analyz th data within sits: Y ij = μ + b j + i + ε ij, whr Y ij = prforman of th ramt of i th lon within j th blok; μ = ovrall man, b j = random fft of th j th blok; i = random fft of th i th lon, E( i ) = 0 and Var ( i ) = σ, ε ij = random rror, qual to th intration btwn th i th lon and j th blok, E (ε ij ) = 0 and Var (ε ij ) = σ. Th analysis of varian was prformd using produrs of th softwar Slgn, vrsion 1.0 (Rsnd & Olivira, 1997) to stimat ovarians. Tsts for normal distribution of singl tr data, lon mans data and rsiduals wr mad using th Gns softwar, Windows vrsion, 001 (Cruz, 001). A fw missing masurmnts in th trials wr du to low dvlopmnt of th trs girth, mainly xplaind by th lak of ompatibility btwn rootstoks and sions. Broad sns hritabilitis on plot and lon man lvls wr alulatd. Following Cottrill Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

48 Gonçalvs t al. (1987), a distintion was mad btwn diffrnt hritabilitis dpnding on th purpos. Th alulatd broad sns hritability on plot man lvl is, in this as, usd to alulat gnti gains by mass sltion on sits similar to th trials, whr th blok ffts an not b takn into aount. Th broad sns hritability on a lonal man basis is appropriat forditing gnti gains from sltion among th lons in th trials. In this as, th fft of bloks an b aountd forior to sltion. Th plot man broad sns hritability h p was thrfor alulatd as: hp = σ /( σ + σ b + σ ), whr σ = gnti varian or ovarian omponnts btwn lons, σ b is th blok varian, and σ th rror varian omponnts. Th broad sns hritability on lon man lvl h m was alulatd as: hm = σ /( σ + σ / b), whr b is th harmoni man of ramts pr lon in th trial. Approximat standard dviations of hritabilitis wr found using th dlta thniqu (Bulmr, 1980). Th gnti orrlations btwn trait x and y within sits wr alulatd aording to Falonr (1989) as: rg = Cov /( σ x σ y ). Th gnti ovarian Cov was obtaind by using man ross produts from th Gns produr. Approximat standard dviations of th gnti orrlations within sits wr alulatd aording to Falonr (1989). Th phnotypi orrlations r p btwn trait x and wr alulatd as th Parson orrlation using th produrs of th Sanst omputrogram (Zonta & Mahado, 199), sin th phnotypi orrlations in this as inludd all lmnts of ovarians/ varians: th gnti, th blok and th rror ovarian/varian. So, aording to Falonr (1989):, whr Cov is gnti ovarian btwn trait x and y; Covb is blok ovarian btwn trait x and y; Cov is th rror ovarian btwn trait x and y; σ and σ b ar lon and blok varians. σ subsripts rfr to traits x or y. Statistial analyss aross sits Th linar modl for th analysis of varian aross sits had th following omposition: Y ijk = μ + i + b j(i) + k + ik + ε ijk, whr Y ijk is th prforman of th ramt of th k th lon in j th blok within i th sit; μ is th ovrall man; i is th fixd fft of i th sit; b j(i) is th fixd fft of j th blok within i th sits; k is th random fft of k th lon with E( k ) = 0 and Var ( k ) = σ ; ik is th random fft of intration btwn i th sit and k th lon, with E( ik ) 0 and Var( ik ) = σ ; and ε ijk is th random rror du to lon by blok intration within th i th sit with E(ε ijk ) = 0 and Var(ε ijk ) = σ. To b valid, th tst of gnotyp-nvironmnt intration in th modl rquirs: 1) homognous lonal varians aross th diffrnt sits to avoid a gnration of gnotyp-nvironmnt intration du to sal ffts; ) homognous rsidual varian among nvironmnts; and 3) normal distribution of th rsiduals (Burdon, 1977). Ths onditions for th modl wr fulfilld, tsting th gnotyp-nvironmnt intration onrning numbr of latx vssls btwn th four trials. Howvr, som transformation of data from trial D, was nssary in th analyss btwn trial A- D, B-D and C-D. Th transformations implid a rdution of th valus by a fator qual to th ratio of varian of rsiduals from th two sits in th analyss, to assur homognous rsidual varians aross sits. In thos ass, it was latr nssary to us logarithmi valus to gt normal distributions of th rsiduals. Gnti orrlations aross sits for all traits wr alulatd using typ B ovarians (Burdon, 1977), and th gnti varians of th traits within sits, onsidring th sam trait masurd on diffrnt sits as diffrnt traits. Thrfor, th gnti orrlation aross sits aording to th dfinition in Falonr (1989) was: rg = Cov g /( σ ) g σ x g. y Corrlations aross sits for total numbr of latx vssls wr thrfor onsidrd as intra lass orrlations on basis of th varian omponnts in th modl usd for th analyss aross sits. Th intra lass orrlations is altrnativly alulatd as: rg = σ ' /( σ + σ ), whr is th orrlation aross sits; σ is lon varian aross two ' sits; σ is lon-nvironmnt intration, adjustd for sit diffrns in varian, stimatd as: ' σ = σ ( σ x σ y ) /, whr σ x = gnti or lon varian on sit x, and σ y = gnti or lon varian on sit y. Eisn (1994) found this mthod to giv biasd stimats in as of unbaland data, unlss gnti varians and rsidual varians on diffrnt sits ar qual to ah othr. Evn though data wr not fully baland, givn that gnti varians and that rsidual varians on at last thr of th sits (A, B and C) wr similar, th formula dsribd by Frnando t al. (1984) was onsidrd appropriat to stimat th gnti orrlations, alulatd as: y1 l1 y = = y l μ1 Z1 Z1 0 + ug + u I + μ Z Z 0 whr y i is th n j 1 vtor of data for trait i, i -= 1, and n j is th numbr of obsrvations for trait i; μ i is th xptd valu of trait i and l j is an n j 1 vtor of ons; u G is a vtor of avrag gnti-group ffts Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

Girth and rubbr yild in rubbr trs 49 of nvironmnts; u I is a vtor of gnotyp x nvironmnt intration ffts; and i is a vtor of rsidual ffts. Diffrns btwn lon mans on on sit and lon mans on anothr sit wr alulatd to invstigat diffrns btwn lons, onrning snsitivity to hangs in th nvironmnt. Th diffrns wr found aftr adjustmnts du to diffrns in varians btwn two sits, x and y. This was don by multiplying th lon man on on of th sits with rgrssion offiints b yx. Th diffrn btwn a lon man on sit x and y, dif was.g. found as: dif = ( byxmx ) m y, whr b yx is rgrssion offiint to adjust for diffrns in varians btwn lon mans on two sits x and y, in this as usd to mak mans on sit x omparabl to mans on sit y; mx is lon man on sit x, and m y is lon man on sit y. Th rgrssion offiint b an b found as: b yx = Cov / Vx = ( r Vx Vy ) / Vx, whr Cov is ovarian btwn lon man on sit x and sit y, assuming zro gnotyp-nvironmnt intration; r is th lonal man orrlations aross sits, qual to 1 sin th gnotyp-nvironmnt intration is assumd to b zro; V x is th lon man varian on sit x; and V y is th lon man varian on sit y. Diffrns wr alulatd onsidring on sit that was both indpndnt and dpndnt in rlation to anothr sit. An analysis of varian of th numri diffrns btwn lon mans on diffrnt sits was thn arrid out to xamin if som lons, in gnral, rat mor snsitivly to nvironmntal hangs than othrs. RESULTS AND DISCUSSION Girth at panl opning Diffrns btwn lons wr highly signifiant in A and B trials, and individual broad sns hritabilitis wr about 0.55 in trials A and B, and lowr ( h i = 0.47) in trial C, whih probably an b attributd to a sampling rror du to missing trs in th trial, and variabl growth onditions within th rpliation (Tabl 1). Th phnotypi standard dviations wr btwn.4 and 3.8 m, and th avrag of girth at panl opning in th trials was about 4.9 m. Compard to othr traits, hritabilitis wr quit low but absolut valus of th phnotypi standard dviations Tabl 1 - Varian omponnts and broad sns hritabilitis on th diffrnt sits. Furthrmor prdit gains (G) by phnotypi sltions with sltion intnsity i= 1.35 and by sltion among th lons in th trial, using th lonal man hritability and with a sltion intnsity of i= 0.81. Data from sixty svn 10 yars old lons of Hva, tstd at fiv loations in São Paulo Stat, Brazil. 1 Traits Sits 3 Varian omponnts CV% x 4 Hritabiliti s Phnotypi Sltion sltion in trial σ b σ σ σ p h p s.. h x G G% G G% Go A 0.1695 3.1096.7888 6.0673 3.9086 45.1161 0.515 0.16 0.7699 0.699 1.53 0.9710.16 B 0.0768 8.344 5.8644 14.836 7.5969 38.0195 0.5840 0.09 0.8101 0.7895.08 0.575 1.51 C 0.3763 1.1790ns 5.6887 6.9440 3.0705 35.3633 0.4697 0.10 0.3834 0.594 1.50 0.8434.38 D - 6.977ns 13.5096 19.8073 5.76 48.005 0.3179 0.08 0.5831 0.498 0.09 0.888 0.60 E 0.8493 7.0146ns.771 10.6357 5.505 48.1567 0.6595 0.09 0.88 0.8916 1.85 0.9655.00 Yr A 0.198 9.8353 7.4995 100.5546 18.0751 53.3061 0.93 0.9738 0.48 0.465 0.8179 1.53 B - 18.0633 11.3164 9.3797 18.6175.886 0.6148 0.87 0.831 3.64 0.8001 3.50 C 0.443 18.6988.5188 1.09 14.9008 9.000 0.863 0.05 0.9570 1.1670 4.0 0.7580.61 D - 133.1347 16.864 149.411 9.0448 39.761 0.5910 0.03 0.9608 1.046 3.03 0.7893 1.99 Bt A - 0.1001 0.0396 0.1397 5.5398 5.7105 0.7165 0.8834 0.9687 16.96 0.8058 14.11 B - 0.1117 0.0333 0.50 5.8980 4.319 0.6787 0.03 0.3950 1.058 4.87 0.8100 19.14 E 0.0003 0.1876 0.061 0.140 7.9069 5.585 0.87 0.9535 1.185 1.86 0.8165 14.60 Lv A - 0.557 0.301 0.8713 5.5719 13.348 0.636 0.03 0.8378 0.8553 6.4 0.805 6.0 B 0.0577 0.3444 0.1397 0.5418 7.6517 7.6700 0.6356 0.8809 0.8593 11.0 0.8061 10.51 D - 0.4010ns 1.157 1.5537 4.9361 1.880 0.580 0.05 0.5106 0.3488.7 0.736 5.74 E - 0.390ns 1.1956 1.4946 3.7069 14.7514 0.000 0.06 0.486 0.704 1.83 0.89 4.53 1 Go = girth at panl opning, Yr = rubbr yild, Bt = bark thiknss, Lv = total numbr of latx vssl rings. Jaú (A), Mooa (B), Ribirão Prto (C), Votuporanga (D) and Matão (E). 3 σ b = blok varian; σ = gnti varian btwn lons; σ = rror varian; σ p = phnotypi varian. 4 h p = broad sns hritability on plot man lvl; h x = broad sns hritability basd on lon mans. P < 0.01; ns = not signifiant. Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

50 Gonçalvs t al. wr high. A modrat sltion intnsity i = 0.8 qual to a sltion of th 43% bst lons will giv a prditd gnti gain about 0. m whil a mass sltion outsid th trials will giv a prditd gain about 0.73 m with a sltion intnsity i = 1.35, qual to a sltd proportion about 14%. Th man girth of about 43 m on panl opning in th trials agrs with formr findings by Gonçalvs t al. (1998). Invstigations of immatur rubbr tr in Amazon rgistrd individual broad sns hritabilitis of about h i = 0.57 (Gonçalvs t al., 1983). Hritabilitis found in this invstigation sm small ompard to that. Mor pris masurmnts.g. on adult trs, would probably hav ld to smallr rsidual varians and hn highr broad sns hritabilitis. Th phnotypi standard dviations wr similar to thos rportd by Gottardi t al. (1995) foirth at panl opning. Rubbr yild, bark thiknss and total numbr of latx vssls rings Clonal diffrns in rubbr yild and bark thiknss wr highly signifiant on all sits (Tabl 1), with modrat to high individual broad sns hritabilitis, ranging from 0.59 to 0.9 for rubbr yild and from 0.68 to 0.87 for bark thiknss at svn yars. Prditd gnti gain for bark thiknss was.41% for a sltion intnsity of 50% basd on lonal mans. Th prditd gain from a mass sltion is about 0.60 to 19.14% for a sltion intnsity of 14% (i = 1.35). Th hritabilitis on lon man lvls and phnotypi standard dviations ar los to thos rportd by Gonçalvs t al. (004) in th sam and othr 10 yar old trials. Hritabilitis for rubbr yild and bark thiknss on sit D wr lowr ompard to th othr sits, probably as a rsult of vry unvn growth onditions. Total numbr of latx vssls rings had a modrat to low broad sns hritability on individual tr lvl at 0.43, and a offiint of varian at 5.46%. Prditd gains ar about 6.70% for a sltion of th bst 50% lons in th trials, and about 5.54% following mass sltion of th 14% bst individuals. Corrlations btwn traits within sits Girth at panl opning showd in gnral vry larg gnti and phnotypi orrlations with bark thiknss (Tabl ). This is in agrmnt with findings of Gonçalvs t al. (1989) for 33 IAC lons, and Gottardi t al. (1995) for 11 lons at th fourth yar of yild. Also Gonçalvs (198) found, in wild mothr trs from 10 sits, pronound diffrns in bark thiknss btwn fast growing and slow growing trs in th Amazon vally basin. Howvr, fast growing trs of Hva hav a tndny to maintain a highr lvl of positiv girth orrlation with bark thiknss. Gonçalvs (198) found largr standard dviations within th groups, whih would suggst a strong gnti influn on bark thiknss. Rsults from Ho (1976), who invstigatd 4 Hva lons from diffrnt trials, showd that th fastst growing trs also had a gratr bark thiknss ompard to th slow growing groups. Howvr ths rsults should b intrprtd with aution givn th rdud numbr of trs in ah group. Th stands wr furthr haratrizd by high sit indis, whih might xplain why th orrlation with growth for th trs in th most fast growing group was apparnt in th bark thiknss. Morti t al. (1994) and Book t al. (1995) working with prognis in juvnil stag, also found that fastst growing plants tnd to maintain high lvl of bark thiknss. Vry high phnotypi and gnti orrlations wr found btwn girths at panl opning and rubbr yild (Tabl ). Th high positiv orrlations found btwn rubbr yild and numbr of latx vssls is in agrmnt with formr rsarh findings (Ho, 1976; Tan, 1987; Narayanan t al., 1973; Gonçalvs t al., 1989; Kalil Filho, 198; Liy & Prmakumari, 1988). Modrat to high positiv phnotypi and gnti orrlations btwn bark thiknss and total numbr of latx vssls wr also found in th trials (Tabl ). Similar rsults hav bn found by Gonçalvs t al. (1998) in four trials. Effts of sits and gnotyp-sit intration Girth at panl opning - Although diffrns of girth at panl opning btwn th sits wr small, thy wr statistially diffrnt, xpt btwn trial A and B (Tabl 3). Vry high gnti orrlations aross sits showd that girth at panl opning has vry littl gnotyp-nvironmnt intration (Tabl 4). Howvr, all hypothsis of gnotyp-nvironmnt intrations qual to zro btwn diffrnt nvironmnts wr rjtd in th analysis of varian aross sits. Th analysis of varian of th diffrns btwn lon mans on diffrnt sits rvald signifiant diffrns among lons onrning snsitivity to hangs in th nvironmnt (Tabl 5). Rubbr yild - Signifiant diffrns btwn th sits wr prsnt Sit A had th highst rubbr yild and sit B th poorst rubbr yild. Gnotypi and phnotypi orrlations for rubbr yild aross sits (Tabl 4) ar vry high, ranging from 0.63 to 0.89 (phnotypi) and from 0.64 to 0.9 (gnotypi), agring with prvious stimats by Gonçalvs t al. (1998). Th analysis of varian btwn lon mans of rubbr yild on diffrnt sits showd that som lons, in gnral, wr mor snsitiv to hangs in th nvironmnts (Tabl 5). Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

Girth and rubbr yild in rubbr trs 51 Tabl - Phnotypi ( ) and gnotypi ( ) orrlations within Jaú (A), Mooa (B), Ribirão Prto (C), Votuporanga (D) Matão (E) sits among girth at panl opning (Go), rubbr yild (Yr), bark thiknss (Bt) and numbr latx vssl rings (Lv) of sixty svn 10 yars old lons of Hva, tstd at fiv loations in São Paulo Stat, Brazil. Sit/traits A - Indiana Corrlation Typ Rubbr yild (Yr) 0.5847 P < 0.05; P < 0.01; ns = not signifiant. Traits G irth at panl opning (Go) R ubbr yild (Yr) thiknss (Bt) 0.677 thiknss (Bt) 0.799-0.1333ns 0.6968-0.1547ns Latx vssl rings (Lv) - 0.357ns 0.7615 0.7458 B - Mooa Rubbr yild (Yr) 0.6510 0.73-0.ns 0.6493 0.7593 thiknss (Bt) 0.6461 0.688ns 0.7039 0.1708ns Latx vssl rings (Lv) - ns 0.8007 0.679-0.141ns 0.938 0.971 C - Ribirão Prto Rubbr yild (Yr) 0.735 0.638 thiknss (Bt) 0.6154 0.06ns 0.651 0.106ns Latx vssl rings (Lv) 0.033ns 0.897 0.9075 D - Votuporanga Rubbr yild (Yr) 0.7436 0.650 0.151ns 0.631 0.853 thiknss (Bt) 0.7781 0.3841ns 0.865 0.351ns Latx vssl rings (Lv) - 0.619ns 0.5944 0.5913 0.615ns 0.8738 0.815 E - Matão thiknss (Bt) 0.7563-0.814 - Latx vssl rings (Lv) 0.0865ns - 0.5537 0.1415ns - 0.5771 Corrlations btwn diffrnt traits aross sits Gnti orrlations btwn diffrnt traits aross sits, using th B ovarians btwn lon mans as dsribd by Burdon (1977) wr in gnral smallr and of opposit signs ompard with th orrlations found within sits, although diffrns wr small (Tabl 6). Probably th diffrns asrib to influn of sampling rrors, gnotyp-nvironmnts intrations for th traits, and by th fat that nvironmntal orrlations ar zro aross sits, but ngativ within sits. Th modrat gnti orrlation btwn rubbr yild and bark thiknss in trial C almost disappar btwn rubbr yild in trial C and bark thiknss in trial A, as wll as th strong orrlation btwn girth at panl opning and rubbr yild in trial C disappar btwn girth at panl opning in trial A Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

5 Gonçalvs t al. Tabl 3 - Rsults from th analysis of varian aross sits onrning girth at panl opning of sixty svn 10 yars old Hva lons, tstd at fiv loations in São Paulo Stat, Brazil. Trials Sour D.F. A-B A-C Bloks/sits 4.044 17.1061 0.903 0.4 19.0974 Sits 1 6.5447ns 100.6571 86.5558 14.7048 158.7315 9.5077 1.896 13.54 5.538 7.43 x sits 4.60ns 6.999ns 5.439ns 11.5837ns 9.7507ns 134 3.0170 4.391 3.073 5.4868 4.640 T rials D.F. B-D B-E C-D C-E D-E Sour Man Squars Bloks/sits 4.8945 4.7589.0199 15.7839.176 Sits 1 1584.49 967.7760 740 34.686 75.5761 19.0040 3.3108ns 37.3913 46.4386 6.0787ns C lons sits 8.1397ns.0311 4.743ns 13.893ns 4.408 134 3.4573 5.8717 4.6793 7.093 5.971 P < 0.05; P < 0.01; ns = not signifiant. Sits: Jaú (A), Mooa (B), Ribrião Prto (C), Votuporanga (D) and Matão (E). A-D Man Squars Tabl 4 - Phnotypi ( ) and gnotypi ( ) orrlations aross Jaú (A), Mooa (B), Ribirão Prto (C), Votuporanga (D) and Matão (E) sits foirth at panl opning (Go), rubbr yild (Yr), bark thiknss (Bt) and latx vssl rings (Lv) for sixty svn 10-yar old Hva lons in São Paulo Stat, Brazil. A-E B-C Traits GoA YrA BtA LvA GoB YrB BtB LvB GoC YrC GoD YrD LvD GoB 0.6456 0.3489ns -0.1999ns 0.0039ns 0.7745 0.359ns -0.380ns -0.033ns YrB 0.5038 0.8005 0.108ns 0.3159ns 0.4791 0.8485 0.0740ns 0.3036ns BtB -48ns 0.30ns 0.7017 0.7519 rg - 0.644ns 0.5875 1.118ns 1.4148ns LvB 0.4465 0.746 0.0148ns 0.316ns 0.488 0.7557 0.0099ns 0.3605ns GoC 0.8469 0.46ns -0.5041-0.3753ns 0.5103 0.1700ns -0.413ns 0.1651ns 0.6819 0.703-0.856-0.4474 0.558 0.6199-0.836 0.1646ns YrC 0.045ns 0.6363 0.38ns 0.7606 0.35ns 0.7445 0.349 0.5515 rg 0.060ns 0.649 0.3493ns 0.8374 0.3471ns 0.7915 1.390ns 0.5640 GoD 0.5419 0.4610 0.1680ns 0.156ns 0.61 0.76ns -0.0643ns - 0.0733ns 0.6061 0.3500ns 0.8886 0.5857 0.3109ns 0.990ns 0.5911 0.3013ns -0.308ns - 0.0574ns 0.898 0.5401 YrD 0.60 0.8865 0.0199ns 0.586ns 0.4344 0.7995 0.3136ns 0.5004 0.4996 0.7416 0.703 0.95 7ns 0.307ns 0.5313 0.9361 0.544 0.546 0.8500 0.7907 LvD 0.4138ns 0.5004 0.5641 0.3786-0.0863ns 0.3368ns 0.0900ns 0.1996ns 0.513ns 0.333ns 0.760 0.6843 0.909 0.7560ns -0.971ns 0.371ns 0.088 0.3173ns 0.8397 0.4639 GoE 0.70 0.581 08ns -0.071ns 0.5516 s 0.716ns 0.0379ns 0.530ns 0.5361-0.5095 0.6190 0.5095 0.7919 0.894 0.64 0.0145ns - 0.830ns 0.614 0.449ns 0.013ns 0.645ns 0.77-0.0773ns 0.8315 0.5496 1.1864ns BtE 0.001ns 0.0683ns 0.7631 0.4136ns 0.0140ns 0.193ns 0.3415 0.35ns -0.110ns 95ns 0.1571ns 0.049ns 0.6869 rg 0.74ns 0.0816ns 0.8 0.456ns 10ns 0.33ns 0.1446ns 0.3968ns -0.0671ns 0.0646ns 0.056ns 0.0354ns 1.0077ns LvE -0.593-0.1393ns 0.7051 0.4795-0.795-0.80ns 0.3671ns - 0.81ns -0.6858 0.059ns -6ns -0.1875ns 0.448ns -0.9769-0.1375ns 0.1795ns 0.796-1.1535ns -0.45ns 1.474ns -0.1144ns -1.4191ns 0.1834ns 0.1780ns - 0.3630ns 1.598 P < 0.05; P < 0.01; ns = not signifiant. Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

Girth and rubbr yild in rubbr trs 53 Tabl 5 - Rsults of man squars from th analyss of varian on diffrns btwn lon mans on diffrnt sits for girth at panl opning and rubbr yild of sixty svn 10 yars old Hva lons, tstd at fiv loations in São Paulo Stat, Brazil. S our D.F. P < 0.05; P < 0.01 Girth at panl opning MS F valu Clon 6 37 6.8 0.0001 7 48 - - S our D.F. MS Rubbr yild F valu Clon 6 169.38 0.0001 7 71 - - P> F P > F Tabl 6 - Rsults from th analysis of varian aross sits onrning rubbr yild of sixty svn 10 yars old Hva lons, tstd at fiv loations in São Paulo Stat, Brazil. Trials Sour D.F A-B A-C Man Squars Bloks/sits 4 308.157 77.6571 78.054 Sits 1 3946.3499 318.9030 838.840 69.9573 154.760 386.7198 x sits 68.4 111.9687 16.7319 134 18.014 9.163 14.3810 Trials Sour D. F B-C B-D Man Squars Bloks 4 9.5073 9.906 6.0914 Sits 1 15.0619 146.318 103.5364 171.977 471.9499 35.9145 C lons sits 57.0538 93.8039 11.3910 134 14.109 19.306 9.406 P < 0.05; P < 0.01; ns = not signifiant. Sits: Jaú (A), Mooa (B), Ribirão Prto (C) and Votuporanga (D). A-D C-D and rubbr yild in trial C. Possibly th hang in orrlations aross trials A and C ar ausd by gnotypnvironmnt intration for bark thiknss and rubbr yild, whih liminats th orrlations btwn thos traits and girth at panl opning rubbr yild sn within C. Finally, masurmnts of girth at panl opning at on sit sm to b nough, givn th vry high gnti orrlations aross sits. Considrabl gains in rubbr yild (abov.0%) ar possibl vn with modrat sltion among th lons. Th gnti orrlations aross th sits did not larly indiat any signifiant advantags, using targt nvironmnts. thiknss showd th xptd positiv high orrlation with rubbr yild and th trait must dfinitly b takn into onsidration whn slting for rubbr yild. Hritability and standard dviation for total numbr of latx vssls wr modrat. Th potntial for improving th total numbr of latx vssls is high with modrat hritability and positivly orrlatd with girth at panl opning and rubbr yild. ACKNOWLEDGEMENTS Th authors thank Fundação d Amparo à Psquisa do Estado d São Paulo (FAPESP) and Conslho Naional d Dsnvolvimnto Tnológio (CNPq) for finanial support. REFERENCES BOOCK, M.V.; GONÇALVES, P. d S.; BORTOLETTO, N.; MARTINS, A.L.M. Hrdabilidad, variabilidad gnétia ganhos gnétios para produção aratrs morfológios m progênis jovns d sringuira. Psquisa Agropuária Brasilira, v.30, p.673-681, 1995. BULMER, M.G. Th mathmatial thory of quantitativ gntis. Oxford: Clarndon Prss, 1980. 53p. Si. Agri. (Piraiaba, Braz.), v.63, n.3, p.46-54, May/Jun 006

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