,'

9n tlo rutno ol'

tle, nact, l"ze,llczt*,n rctkl'

)/lt

GE ETIC I'ECHAIIISI,IS OOI{TROIIII{G I]{IIERITAI{CE OFCLCUV RESISTA CEA D OTHER EOO O IC TMtIs

tN Co$tpiutn hhsutumL

MANzooR HussaM,sc. (tron3.) Agri

A thesis submir0ed in partiat fttfilDenr ofrhe rcquirenenrs for the degrEe of

DOCTOR OF PHITOSOPIIY

IN

PINNT AREEDING AI\[D GENETICS

By

FACULTY OF AGRICULTURf,UNIVERSITY OF AGRICULTURf,

FAJSAI,{BAD, PAXISTAN2008

To

Tt Colt ollcr of Exrniurioo!,

Ulirllsity of Agricultule,

!r'e $c Np.rvisory conmitrec ocrti& lhal

,ubmitEd by M$roor EI'd] Regd. No.

$lisfrElory !!d Monrn !d that it bc proc.ls.d

BIlniF(s) for dF award ofih. d€Ec.

SUPERYISORY COMMITTEE

CMIRMAN

MEMBEE

MEMAER

E5{g.l3fil hlv€

#(kr,-

A,!ifAri)

DrtdcdTo

ItlvtwmEmyl-s

-.

AcKT{oWLEDGEMENTS

Au ptuira ad tton r @ lot 'UlniEht, Alhlt, vho b tta ati'e M of all

ld"dt 4wido6 ard *roeledg., Wo CM n. taLh, Ihdght, tol.nte.l tacha amlhelpikg @llqgra withut ,hi.h it w ,np6ti e Jo ne to @mpl.te this wtL I ofertut the t@bl8t thantu lftn the deepst @ of d, hed to tht Hot, Ptophet HeatMunandrd (Pw be upon hih) wro is h. et a tuodal oJsuidarce dnd k Nledse lol

Virt d.ep enotid o! be"Nlace add sratitudq I ofe n, oppreioti@t to ntwrtht srqnibr Dr Fa4n MuhMt! Azha. Prcle$ol add C^altua", Depa hent ofPla"t Brceding aa.! G@ai6, Url|ersity oJ Agridltw, Faisalabad lo his l,titing help,tdgei6 sugaation, sEp to tt p etida@ ard cl6e sqdiior .lunne h. cordt l oJth* irwtiAatiN od in tte F4gntia of hn Mwip.

me autho, is highl, id.bt d to Ptul Dr, Atif Ali Dep4n@t of Plut B@dingard GeEaiq, Uniwsitt 4,ledaltule, Fa&otabadfo/ hh afdtiorate behdiod, wttl.oMak @t! h.lp ren lered durirg tlte tptt of this edirc studr.

I ale ||b, to etq@s nr d..p @ 4 Fatiltd. to Pr4 tu. Statz.l MaqtoodLBNa Depatln q aap Prrtiol@) U"iwsitt of ,IetialnE, Fat:alabad fd hi;

I dan n nt pl@urc to Mif irs.f of thh oppo unity to eq6t n,@"tPli@n ard Prcfottt! obligltio8 b nt .l%t R@ M. &.al lqbol Khan (pide oIpefoM), D,. Muhodda.t 4tq /w. (Ex Ueadt M'!ai$ arudine D'xior) and

"t, ot d @ ag/6 of@u@ gtotp NUt Failalaba.ldpeiallt Mi N@ir /, Na PSA,

Mi Sdlat Mthdnna.l C&4 .4gri., M| Mrhannad tlmn W, ML MuhMnad Alifuh-t ud Mr. Somar M$tala Ldb lterddht Jd theb aluatt i$pinns ond

I Ied m|xetr helpless in sanclittg ptopct wrds to pay bacL to ny larhe adtuthe |'ho @ d bdh now in hz @fl|due to thd,I @h.d at ttis p4iti6, I daho ptuud otnt famil, ^enbe$

|'ho al\|ays iNpned ed er@rrcged ne in @mpletionol tut Ph.D ttudiet. I pr@t n, ,p.ciat thatk' lM the @e 4 kr hqn. b ,t btuthqtard

'tn 6. ,h@ huds altu

' rcireafot n. i, pmreE.

hst tut not lu4I W ital eibu8 vitA de dz.pat we to ny wile. wad dausttq lor their srppon ad pdia.e shich .tuble rc b conpl4e th. sadr. Ma,t ah siE thm a lone a"d hapry lik '.rnq\

Col{rEms

..--)^y4)r " '1 S|UD|ED

s#|1::";::::w-ii+-,-(

CTIAPTEB TITLE PAGE

1

2

2.1.

zz

2.3.

2.4.

2.5.t.

25.2.

2.5.3.

LIST OF TABLES

USI OI NGURFS

LISTOF APPENDICES

ABBREVIATIONS

INTRODUCTION

REVIEW OF UTERATI]RE

Corcept of dbease in platrb

Plant viEs€6 ae pathog4i. aSots ad th€ir Dehanilm of

Virus infetion dd replication in h6t cels

Gdini vitutes - thei. mode of tEsmilsiorr imp&t onyield and inh4rit re studi€r d global b6risViral dis€ag ofottonMech4is of spr€d of cotbn l@I (l1Dple viruImpet of ctauD on pl.ant Ptodudio d tlobal ba3i3

History of developmmt of qruD in Pakbtan

SymptdDloSy of leal cul and (tuple viral dis€as in

Ve.loF for ttE trMis6ion of qrcuD and aliehale h6is

D*elopmt of CLrUD Ei5t nt vdietig on global bsit

Genetic studie. of cotton le.I @l vins dis* (Cl,CuD).

Mol<rlar studig for @ttm laldl vitu die*IntEitac ol atMdic ard fiber quality Fails, dd tneu

ll't2

14

'14

16

17

2.5.6.

2.5.7.

2.5.E.

25.

18

m

AB25

pGsible asso.btim

CHAPTER TITLE PACE

3.0

3.1

3.1.1

3.1.2

3.13

3.2

33.

3.4

3.5

MATERIAIS AND MBTHODS

ScEning of cott6 templasm aSaiGt cottdt lef tuleitu disease (clcuD)Assdnent of cotton gemplalm in 6eld conditi@Asment of cottd g(fipl6lm udq ELss h@e

Assg6|'qt of otton gerpl.m thrcuA galing

Development of plart Mterial fd inhditarre of CLCuD

A556!tqt of guo9?ic 6polg to CL,CUD in the field

A6shst ot geotypic Epollg b CrcuD 6ds 8la$

Meal,t€mnt of agercmi. and fibr tlait3 in the plant

STATISTICAL ANALYSES

G{eration ne@ dd vdiatue analysis

Btifrtion oI hetero6i! and inbreeding depresion

E6ti@tion oI doninareEsri@6on of tuw 5€@ hditability dd goetic

Componenb of vdide

RESULTS

vdiation in 8do9?ic re6potg to cotton leal tul virud'!€@ (ctluD)Al66nent of six geMation for CLCUD resistafte, ad

Gendation @@ dd variare aElysis

33

33

35

36

37

39

3.5

3.7

3.8

3.9

3.10

ltal

41

43

)A

14

3-11

3.72

4.1

45

t*l

47

4.3.

OIATTSR TTTLB PAGB

Bti@tid of hehr.6i!, inbrceding d?'6i6 and poterr€

Componanb of vdiatio4 6timt6 of na!'w ea$e

h6it blity and 8&elic advarre

DISCUASION

SI]MMARY

IXTEIATURA CTTED

APPEI\IDTCES

45

4b

6

71

7a

g7

92

LI5T oF TABLES

T.ble P.ge

3-0

!.2

4.4

4.4.1

4.5

Cotn \ (Cacrpiun hitutun L.) Sdotyp6 ieted aganltt CtcuD

Cotton leaf @rl vins dise (CLCUD) nting *ale

C@rrents tor partibming oI 6M ol sgt|.€ donS 3ix

teEaiioN inio orth'rAonal comparisds (utde dd HU r97q

Coeffi.ienb of g{eflc effets for weighted le6t sque analysigof strration me@ (M.thd and link, 1982).

34

38

41

3.3. 42

Coeffi.ients of geetic vditue @mponenb for ttE weighd €le6t square analysi. of Eendation vdid.es (Mathd and Jinls,1%2).

Comparbon for the respolg of.ott6 vdieties/lineg to cotton 51leaf.ul virur dil@e (crclD)

Analvsit of vdiee of ratins of 40 cotton veieties/ lic hor 52CLCLiD unds tul<l conditioir

Rdlhg dd srcupins oi (ofion vdieri6/sE iD on rhe ba$ oI 53their r€apqE; to CltuD

Reiffi of.otton gmotypes to CLCUD throu8h anificial 54

Md i-alu6 of v&ious pldt narlt m@ured in the six 55gell@tioN of G6AF u; [dtuE L

Med squares ftom paltitione.l analysis of varid.e plant baits 58,59medued in the 5ix ffneatioa of Ci*ylitn hi@hrn L,

&tiMte of the b6t fit model for gdsation ffi param€tes, 68m6n (m), additiveldl, dmidrelhl, addrive \addiriveIil,additive : domituncE ll, and dmild.e r domiMMe []valq6 for vdioa baib in .otton

Vdidce Comporents, D (additive)/ H (domiMre), F ((Gsprodat of additive and domillfte) 6d E (envircmenr.l)foUminS weidted ar6lt5i5 of cohponsts of vdi@e for

4.7

42

4,7, 69

T.bL D.|.tlpddr Prgr

14,

L9

tuft. f.€drtr qthgd4.dmmdpoEEetu d t r5.ttsq pftyno.tf & dd rvlq|d-"nf lr* 6f rdrq ttdo|vrr trdi|'V d S.tt dcadeaE Ia vdtr brlt hcel

m

Lrsr oF FTGURES

4,1

4,3

4_4

4,6

4.7

4.8

4,9

4.10

4.11

4.12

4.13

4_14

4.15

4.16

Mean valus of qruD .rie.* ssiry of si' cotron getl@tim 56

Mem values for leaf dea (.mr) of six cotton generatioB 56

Md vdu€ for Ftiole length (ch) of six cottm generatiotu 57

M6 valu6lor 3rrpodial l€ngth (m) of sn cott@ g6aation$ 57

Md valu6 for the nMber of bols of six cotton geerarioB 6r

Md valu6fo! boU wetht G) of sr corbon tseratioB 6t

M@ vdue for bol size (.h,) of six cottd generatios 62

Md valu6 f@ *ed.ottonyield G) of six otton gensatim 62

Md values for giming @r tum oI si\ conon g@atiotu 53

Man valu6 for fibq l€ngth (tM) of sr.orbon gedatio6 63

Ma valu6 fd fiber fin€ns (f&/irsh) of six con@ a

M@ valu6 fo fibq @turity (%)of six cotton Sddati@ 65

Men value6 for fiber 6hength (rppsi) of six cotton Sen€rarions 55

Scatt€r diaSra& (c neah l€Iare. (@1and Ftiole ldsrh (cn) 75or sx .otton generafon3&.tt6 diat@ foi 1116 t@I aE (or) dd bous nmbq ot six 75

S.attq diag@ foi tuan leJ arca (n2) and boU weiSht G) of six 75conon gedahoE

4.17 Scaner dlagtu lor ronteaf xs (mr) tud *ed (onon i€td (d 75ot s cotton sqsahm

4.18 ftatter di!8r; for p€tiote logth (d) and number oi bols orsix Z6cotton g{erarions

4.r9 s(atter dtagm for m@n Ftiole lmEth (d) dd bou weight (8) 76ot sx cotton ceneEtim

4.20 Setter diagr; for tran peuole length (oJ and sd cotton 76yield G) of sn conon sserioE4.21 S..tter diagm for m@ total Nmber of bolk dd boll reisht 76G) of sir @non meratioE

P.ge

4.2.

1.8

421

ftatter dl,agtu fordrern 8in tm our (*) dd CtCuD of sir V

S..tt@ diagb!| for !|s flie finerEs (|r&/iBh) atd mturit, Z(%) of sir cotlon get]eiioiEsc.tts diaSid for tlldfilc ffm@ (p&/iMh) and CrcuD of 77

Scatte! diaSm fo! m6 nur$€r of bolh dd CtcuD ofsix n

Lrsr oF AppE DrcEs

D.qldm P.9.Van res of vdiru plsnt Faits msxld in tfE sir 179tor4tjora o{ Go.rlprild tirlllr.r L

II Simple @delation.o.tricidb betvEn v.ios rraib o,f si,AeMatiota of cossypiqfr hiriutunL

tm

TI Meteriologtc.l data 2qB, dder gts house md field 7Aoditi@ !t (FaieLbad) patisr.r

G

ABBREVr rror{s

Ihoosnl Poulds Pa Sc|e l!.n

Cmd[ Dmedic PrDdwt

Cotrd tdf dl viro! di!@

Cotlon R*aEh Stalion Multi4 PatjslEn

Cotnl Cotlon R€S.NI Iolritrt , Mut&4 p|listd

Atub Atri. R64!h InEtitur., Fsisolrb.4 pddsr.!

Neldh6'tut for Agd. ed BioLoet, F.i!.ti!.4 p&isl&

CIP\AD & MdFeltiq, Flue

CDP

CIrUD

HVI

cRs

CCRI

AAIII

NIAB

CIR D

INTRoDUCNO

Cotlon plst ba bd gDM in Eopicd and dlbiiopic.l .limlt€s of lhc eddri@ pt hisro.ic tim6, dd mw i is bcilg dltivrtcd qcnsiv.ly ir t Epdr. climat6 swell (Saud, 193 &d !@ivd dr dl, 199). Narurc h6 favorcd cotion fibd virb ils

miqE (inGm.l) 6!u.rd ald hid-ld6l qu.lity .h@r€rturics which d4lop s .Bult of lhe ints4tion bclwa the Aa€tic dd ovibm€nlsl feroB (Bdmarhy ddPttil, 20Ol: &d Basl, 2003), Of rtc 52 spei6 of @tro4 otrty fou F€ia edoo.sricded &d prcdE. Ainn bl! 6brc c.ll€d lint, m6e fou spai6 t c*stpiun.fttah &d Adtwi.h db.iw, cdt.il 'Old Wold Cottotr" *hicn e ..tiv6 toSoultr@ A!i& md Anic.j ed ihc orhd two @ Corqpr,r iri^utun 6d CosrpiM,arrad"6,, Ie.d s "l\la Wdld Coiroq.. !trd l[6. e n tive ro C.arzt .d NodtSoutn Andi@ (Endrizi e, zl., l98s).

CottD (G6wiun hitutu L) pLrr is e ibporr.Dt nallnt sr. of fhd,which is b€ing u.d in UF world r.xlite i!d6ny, In addition io sneeft. (So.charun

olici'@tuh), ti.. lo,rzA sotiw) tA Mi@ (Za nzy L), c<,/lon belonains ro ,rrd!,rp i5 abo d ioporr!ft c.sh Np of Patist&, In pskisrd! ihc mrion cbD ecours for10.5 % ofthe vatu. dd.d in rgd.ulhft &d .!our 2,4 % to cDp. Thc crcp i! g|om on6 ext Niv€ @a jn ihc @hon b.li. IlE cop ws SroM on 3,21 miUjm hetes ddi.tl€d 14.62 nilli@ bdd of lim duitr8 rhe yo 2004-2005 (E@nonic sery oflatbtan, 2004). Ai p@.nr, conon pldt f..& 426 r.xrit. ni s, 98tJ slindlc ujls. &dI l:O SiEilg f..rdi6 i! P.ti$.! (&hr rd Khd. 2OO2). In a<tdirio4 ro Detitrg rh.n*ds of th€ l@al tqtil. indurry in ih. coulry, th. cbp f€tcb$ norc rhe 60 % of drc

rol3l foEign dch{ge @ir8r @udv rtDud rbc qpo ofw @non,ql fbdnad.prcdu.h, Cotroneed oil, s inDonsl by-pbduct of@tion pldl @ntdbur6 ?5-80 Zo lotot l v.g.{iblc oil obLin d 8Dm oit dd M{il c&?6 i! rh. @uEy (Aliit'r d dr,2002a), ConoEed c.ke, soihd by-pFdu.i. is ! valublc sui.e ofprcrein for onl€. Iri! tlpon d th,r lO % @urry's Lbou fo@ b mpto,€d i! @non pFdudiotr @tkh

.q!||t9.1.... ..................... !4.9*s!.

ptE*ir8 dlt! ad .lli.d ildusti6 (Alhtt d z/., 2002.). Patistd ruls 4d in @ooo

@a ed pbduction dd li s d cxpona of nw colton in the wdld (Cofon R.vi@,

PCCq MIMAI" tul, 200q, Tnc Pujrb k dc l.rditra cotton gtowing pDvi&.

dtibuling !l@ lhd 80 % lo ih. iobl @tion produtid oflatil!! (Mateood, 1999).

Sisrli6cel (Srdual) imprcvdot in cotton @ atd podetion i6 dtribuled 10 favonble

watbq, pbDorg of hi8! yr.lditr8 v{i.t6 dd adopti@ of inPbYed rgicultEal

Aicr the onon lad is pbrt.4 its grc*lh ard dd.lopnmt it $bjel ro veioa

6vim6d cG, i,c, ltigi tmpanft, eiidty, nobr@ sir65 ed dl4k of D6l eddis66 which .flet ui1 aH productior Mtjd cud of low l€ld in @l1on eroM in

P.ti!ta! s. lle arr&t of itMt p6t aDd di!..s ar veioE pldt ddcloPMt srgFs

(Cotld: Rdid of Wodd Situtioo,ICAC,2o0l). Ih. viul dis€l$ cNe no(! dlmag€

the oit6 drs!€s, and i! s@ c.&s,lh. otir dopp.d .ru it ddataLd. At Pcatt!@ thr 20 viEl d!!..s ol coe.n hld b@ d!.cdb.d ir tlF 'cofton Disag

Compddim" oI Ahcdce Pht'lopsholoSical Socidy, bul ofin€s onlv Gdioi viru6

N of @mnic inlonee (8Dm .!d N.l5oq l9E4; Malrld ., al' 1993; dd

NrA.a et .1., 191). h lhe @at pss! tlE widc spF.d of @t!o! lsf .ul litu(Ctcuv) tri|lmjfed by wni. ny , Beni.ir tdbad Gtu (MMr ., al.' 193; H@ed

a al..1994t d Alntt .t a1.,2m2.. b) ha cu]r€d h ojd otlot lo6s in P.ki3t r

Duing 19!92, Paldshn pNdwe{t 12 8 milion btl6 dd r&l@d 3d in ih. world

aad chin ed U,S.A., .nd ws 3@rd to @ft @tbn dPonina.oulri6 Th.@na'

U. 6non pDducli@ ddline4 .nd in 1998-9, lle @utry Prcdlc.d onlv 8 8 nilimb.l€s. Th. m.jor cuc ofthe i.ld d..line b cohon prcdwtion ws thc d@uP of cotton

lcrf cEl viu dis.e (ClCuD), rhich ded.lld the @ufvt dmDv ror lmv ttG

b P.Linar. th. CrCuD ws 6Fl mt d on f.w individud pldl! in Mul14

(Pujrb) in 196? (sG.in ed Ali, 195). Th. dis€& FsisrEd in rh. 6.ld! .r ldirtaEity slil 19s7. h 1989 and 1990, lh. iftidm. of CtauD ift@!.d couiddeblv

srd yicld lo!s6 vdi.d 6@ 22.3 to 68.5 ',4 i! 6mc .@ of Pujtb dc?qdilg d tho

vEieti*, iim. of infetid dd dvircmmtal dditioD (sussain ./ 41, l9l) In lhe

SidlL prevh@ th€ dis.e vas 6$t lwdcd i! 1996, dd hd sift. bd .pidoic in

Norihm Siftlh, tn additiorl ilr€ dise ha alio bco rcpon.d in lhe Balochjst n

pmvirc. &d Nonh W6ttu Fontid povinc. (N]i/FP), Th. deva|ling etrdL of ilisvinl di#c on cotton hav€ .lso b6 dolMdicd in SudE (l&is, l99o).

R.s.eh work6 hav€ nrdc efotu to iddiry fie c&s ol ihe disd., edFport d 1h!1 lhe 6lton ft! wlndalnity lo Cr-CuD &d othq disc.s is du. b tne

rdow 8pn.hc be ofvdi€tiq b.ing plet€d in ih..oftn bel (Iqbal e, al., 1997). The

nmw emctic bal. up of the vdi.ti6 hs 6ulr€d duc lo .xFnsiv. w of 16 difding

taMrs in cDsing prcgtnmd, which uldnat ly bd 1o dintuish 8@tic div.6ity itr

nq smplm (Fouilloq md Buqo! 1988). Itis hqd aDd ini@ting f4ct b4uehoe onvircine wh6 thc msl pndoni@tly 8lrlm culdv6 lilc S-12 &d CIM-240,

which .ovd.d about 44 o/. dd 63 % of th. bral dca, Gpeclivclv, duing 1993,

€xhibiled lh. hi8!6t degr of sw@tibilily lo @don lcaf cul viru' To l&kl€ th€

ptubld of CICID in the tujor @non growins .l@ of Puja!, a nMbd of lt:rcsi6

w@ pu1 in pl&e. I1F8c includ.d cl@ culiivali@ of conon crP usin8 ,mpovcd

sgrononic pr:.tic€n nddatory cort ol of whn ny (1h. v6br of cl-cuD), climiirlon

of iser p.sis uing Fsticidd, tuid bd on the olti!.tio! of suecltibl. v&.lics Ih€

mosl dep.nd.bl. m€surc mphaizcd lo b€ it lh€ ddetopn@l of Yitu cisl&l veieties

io @p. with ilc aiming situlrion of cl-clD in the @uEv Grugi! a al. l99l:' ttd

Ali, l9?). Cl€arly, such a breedinS Pregllmc wolld bc iucestul if the sd.tic bde

of ilE bre.di4 popuhrio! is bo.d.ned (Md.ditb 1991| sd Ali, 199)

To .@ut r thb mo@ of CtCuD @iddiq the avtlabl. cond s.mPlarm

w6 *@n d by eliq res@h6, but nde of ilEn showed resbtancs Th@forc' sme

cxoric 8@type *4 irtmduc.d dwing 1993, &d t6ted ddd ou lalt grewing

.ondilion!. Fa of lh.n rmelv Ccdix, LRAL 166, Iab&.b (69) I I '

Reba P_288 ed C_

l?43 $ov.rt r6isl!@. lo cLCuD ud..lo.al conditioc cedix, . €ultivd toE El

SAlvlrto(, i! tDoM for ii! hieh Gist!re !8.itst @totr lcd cmplc liB (wilsn e, aa

198q. fts @non b@de6 in Pakiste @tcd io utiliz! il| qolic g€nelic @!EG r@t!on ibpbv@61 pre8llmc sd d a t6ult & @y of new @non vdi€tiq nm€lv

CIM-IIOO, CIM435, CIM-441, MNlt-552, CIM-4'16, CIM_443, ClM482' dd FH_634,

FH-9OI ed FH'IOOO waE dd.loP.d ai difi@d 6@h cmlssi dd @odcodoi for

.q!.rg!

gddal cultiv.tion in rujab lroeinc.. A.ll th6. lwly brcd cultil|Ir ibowcd |ts,s1dce

ro CLCuD lo a vdilllc dcg€€, ed th6 coti@ yicld incEs€d in thc @non bcll. Ih.culiE ClMl100, althoue! tdl lrowins {ith dde foliaSE sd l.te in baiuily, wa

!t!d ert Nivcly in hylddizldon pregllm6 by @non bEtdc6 b d4lop bt!€ding

mat dal fo. diesc @ieloc.. nE wida cuhivali@ of th€ Bisldt veieiG i! Dajot

rrowing ajg elghtly ircrGed.ofor producriol How.vB, on @oul of th. €rrEric

8po@ of vs.nd b.rvimMdlal flrctutios @d p@r fibd qu.lty ch@lerntis,

pdiculdly lit r fin€n $, tn€ cultivatior of ltr.e v{i.ti4 ba io b. di$onrinu€d jNt

rn6 r o 2 y.6 of th.ir tfta rcld& Ir&r oa tm hi8! yr.lding edi6i6 @.lvNIAB-999 dd NIAB-I I I Mlh eood 6bd qu.lity dd b.ttd @isldnce b CrcuD ed

hq1 roltuc w@ .ppov.d for gdenl clltiv.lion duing th. yc6 2003 sd 2004,

Concwd, noleuld ttudi* lor i!. ilettifcalion of viEl sf.io cauing

CtfuD w.r. initbled ir the @ul'y. ft. CLCuD w$ foud to b! coled bv . coqldof whileiy.tmnin.d Gdini viM (Md!6i et dl., 1993: dd Hued er ai ' 194),

b.longilg lo Besmovid5.., whi.h she dilldat ldck of iddritv aidr 4h oihd

(ZDu r! dl., 1998; Mallld €! di., 199; s@ ,r al, 2000; ad M?@or et al '

20ots).

Thc @jodty of @nohi4lly importdi Gdini vitu6 f.ll ilto tiie 8du B.gonoYifl$

which Dl66tly G@hpllc cl@ ro | 00 disnrct 3!-i6, ed all of thd @ tE@ined

by wfit€fly (rerirta tard.i Gfl.), dd IobM b inf6t onlv dicotvl€donous Plantr

(Brnldon sd Ms*b4, 1995). Th€ CrcuD b€ils qualrtativ. in lane (spsi6.

ci.tte) is bctl4ld 10 bt eoldn d by mjd s6ea ed thu llers E cl6ti6'd irto

fN w.lld.nn€d non-ovdlalping cl6$, dd ir contEi 8@eral eislmce 's

qEfiativc i! !rr@ i.c. ph@ltT€ 8iv. ti* to !@y p@dv ov.rleping ch5s6. thuih. mmagm.nt of Esislon@ by conbnilg rutipla Bilidce 8@* into a 6it8le pldt

g!rclype (goE prrmidilg) sho* er€dd dwbilitv, be.ue mutalon oa multiple_

virul@ is udikely lo @u it ! psr popuLrion (R!rdh.w., 1999) or i! !.dtogd, Tltc

inlEritqce of h@t-pLdi 6istonc. n4lunims (P@n.4 4l' l96i Hmmond dd

J@6. 1996) is knom for difcrdt trait! @fding @istt&e .Srinsl th. in@t p€i

conpld.s &d dis.4t.

'Ih. infomdion w?ibblc !o ftr clclrly susa$a trl disesc rt3istarc. of the

bosr b @r a fretion of rhe hosr gmtyF doG; n is &t@incd by U. Sdt}! of thc

paihogd a well. Dilce Gbt!rcc of a pldi is ihe dticul.lion ol int@tiu of 86io '.ytopLsic complcx of th€ horl with th. @n6pondinA irEr&tion oI gdj. x

ctropbsic @opld of llc p@iG (Iin! 1989). The typ6 of intltEtio @ ds!trdtcd by the dvibmdt &iing lpon hosi ed pathogd (p.Eird) i.divitually. At

tn bioch@ical lcvcl, the idcrs.bor ilvolv.s e eucilor nolQulc prodrced by the

!@irc lrd . cotrrlDording Fcpto. in tbc tort @sgl, I95i Stttt wi€ d ar. 1995;

&d kd, 1992). Both Gi.r&@ dd avirule* Cpn6 N Ir@lly doniMr nor

(1955, l97t) p@tuirr.d the me.pt of gdaforg@c El.tiotr$ip bctw6 . ho€t ed ispdbogd h6!d on his wort @ lis.Gd (It @ Krarisiad) ru!t. A@idiry ro 0!3

htloihqis, fo. way r6isrul gd€ p6dl in thc ho!t, ll|e pathogcn hs . ga. for

virul@, ed n h&l b6 dt.nd€d sub.quddy to olhd b&l6al ard nlreal

parhctstlEs (Gtdl dd Rotfe" l99oi dd K..a 1992). Sw.4liblc Er.tid culb odv

wbd ih. pathoS@ is .ble 10 nstch all th. Fsislant 846 pGcnt in ihe host wilh

epopridc virulft. g66. If otu d turc sbta gd6 e ml natcb€d bv rh.

Ddlhog@ wth ln. .@oPnd! virultu gd6" GistlM ractior i3 Gultq! l,tq l[c@nccpt of epn -for-gde int€Fti@ h.d bm tully developed in Plet vitud

patl$,5l@ (B&lcomb. .r ai., 195).

To @nb!t thi! siou dr6*e ro @tto! culre wiih ir! bo6r Prcmw.d.leirindal efiets in ?atishn ed Sudd .lfolts m.d. bv lle €dliq lt*dhc6 in

br€.ding CICID si5tlrr pl&t midilt vd.ri6 (Ali, 1997; rtDgin et 4.r., 2001)'

sisnify thc efrdtivm6s of th. oeledoB ed el*tid of individual pldls toD

s%Egsting populalios udd th€ hir! vi6l load Gsbel, 1929; Gila ed Nou, 19691

..d Siddiq, 1968). Obwdios of @lid rq€th eo*6 ir indioting th. non

sc.priblc staeps of @tton pl6t k' CrauD (ErNou sd Abu Sdfi, 1966; El-Nou'

196? rnd Alhra. er 2r.,2004), dGpodmy ofCLcuD for iB te@ord pafteM s!r.&l on

6doB liLq t q.dture, huidity, wror poPdalio!, ptlv.iling vitd load (Gin tldNou. 1969i Khd ,r al l98r Kho dd Kh& , 2000; Alhld et 41, 2003; dd sh'mmd Pirhr, 2OO4), re|. of &rvcn f&tob in thc aPR.d of tropical whitcllv'tresitted

.flr.PlirJ

s€rmivitud of dilfdmi cmps (FuScne e, ar,l993) h divffi oviFm.nls wa al6o

E4on.d ir dly 6ndin83.

Dinirirhing of lpnetic div4ily in n* SmplM s mjor caus of

wlnorlility oa cottor crcp !o CrcuD (Fouilloq and Bm@q l98Ei ]qbrl .l ,r, 199r,

!6?de of Sdclic vdialility for vi.ddc. .nd FAo,& ag.id CrclD om8 thc

spei6 i.e. O, borfuN. antl G.hirsutun lEbbzls,1916), C.hirsutun 8d G.alboran

(!d@ @d Sulto, 2OO5), ad qilhitr lhe tpdi6 (Alh:r et zr', 20O2a) n..ded for lhc

brclddingofgaelic bsc dd dolution ofCt uDeskt nl cotion vdieliB (Ari, 199)

Linitcd s@* to bred plel nlicrial hlving 6iste. lsel a hiehlv ttsiErant or

imDe twel hd tFa rcluted duc to thc !&k of bfdn.lo8 (i) P.ltm of veation in

CtluD G) @npFh@iv€ ass€3ffit of its es€ti. @ftol (iiD cnesiry tm6 or

viM6 (i, l&t of codtu|ency in vinl tudmlalE tdd rclltrvelv ftqu@t inLr-

s!.cific t@mbiBtron ol cdini virle. In ltre @41ve6, it had bsn npod€d lhal

&c CrclD 6istd@ in bodd @tron .ulti6 miSlt hdc broto do*4 si!@

ppiousty eistdl vdi.des P@ tusclptibl. !o CrcuD in the v.lte district of Pakisiad

(M.|Mr e, at,2OO3; ed Manmood., ai,2m3), Tlrs ot'watid tuggsted lh'

p',',lwe of .oDpfq of sraiE of Ctauv in Pajdsl.. s ildicalcd .and bv Zlb\ et aI

(1998)- Hows.., it had b€d sugigslcd thtt thc n@ cu[ivc !.v. 'tmw adiPLbililv !o

diff.mt erowi!8 6ndinoE, ed ths vieldcd lowd tho uinl€lcd Ne?tiblc cultiG

(Briddon a zl, 200q sd RlhlM ., dt, 2001)

PdioB work on lhc inlEitlM of this viEl di!a& is $dtv' {d whal dc€xist in lh€ tii@ne shows ilat CLCUD is conlrol.d bv a oajor 86e (K!igh! 1948),

sinslc 8@ *ith domimr €frdt (Ali, 1999; Ret!@ ,r 41 2002: ard rl.idd, 2002)'

trclv@@t of a mjor dontuut !pn. in @nt ollirg th. rc6iddc€ of le' @rl liru3

don! with rh. involvdenr oI miM (oodifi4) 866 wd s.al€d bv findinss of

siddiq (19?o). On lh. contdy, R !dh.w. (1999) sd Iqbal d ar @003) EPoned $ar

CLCuD w6 coolrollcd bv two doniMt g€nes dd dupli@t dominsl e?'sl!!F,

6p€!v.lr Findi.S: of R.hnd er 4r eOO5) @el.d tt t lhE 966 vd inblvad in

G ,t6/tuu rsisrdEe to CrcuD, rwo for 6istan@ (Xr...dt etl R d@, 4d a

suppr6g of 6isie. (Jatle} o! th. o$d ned, Ahuja.,,r' (2007) r.foned lh'

invotvmat of tso epn6 wirh dlplical€ doiiM! donimt jnnibitory ud dupli'lrc

.q-b.PF.t

ncc8sivc .?idair od rhrc t!o6 vith riplioai. dodilld ?i!r.si3 i! oonroltiryqr\D r.i.6@. ltowffi, rb. 6!ding. of xhe d zl (2007) r!v..t.d 0l|t rddinvc

96. .facL @tib Gd t|@ to Ei.t&c. rhD to 48.ptibility.

Thc infodllio! o rb! irhdirlE of rliltuEl Dblt rdr! of @mnic ilrqEt i!wcU d@mat d. Mey rnrdid 6 rMb6 of bolk, lint !€!€t gc, bott rcidr, 6bq3ta8ib, h!.ri!t l68q |!gL of Euitirg !r.noi, pLnl tcid! 6.uitil8 bruh nMba&d p@ilid of 6tn Auitiry hdnch, dDwd ncs.livc o! pGitirc .ro!8 .!ei.ti@(Cl6 rld Zte, l9l). Tb..t!o8 rcitirc rdrlid.bip of vr.iou yi.lrd coryonats on

.c.d 6!u i.ld h.lpd bGdd! ro &ld l[G ddiEd g@oF oou.rat, 1992).

Strti6 ty ,i.ry d dr. (2000) Evalcd ih. ol. ot rllcl6 eirh D6itiE lrd!.g.tirc crfdl!! &!d 6.lcctd plet! h.viq le.f.iz. .d !tq. eirlblc for tsii.ut&Fldurid r&.- ft. ol. of .ddiiilc s6cric lFccr i! 6!!lilg tt r..d @trd ictdbd v{iou icld mponest lit !mb- of boll' Dotl wcig li p@ng. brd t cd

l4odd (Yd!v lrd Y.drv., 198?i l(|lry.nd O..& l96E: Wes lDd P.4 l99t;Clranbre d al, 2@!i Mar d .1,, 20o3ri .rd [t6!96d! !!d B.i& 2003), lrd r!.inponlEc of @{dditiv. ga.!.tre& *@ ot 4rd by R.hr. d.t 093t Mcn

daL (2003tI.rd Mci-aa a aL, (2005).

ht qrc of ddiriv. gaaic cfc.rr i! @drelirS th. inh.rirec. of fb6 rr.iklrc rglc lagtt, ft@eq nt!|.iry Dd lradl {4 Efond by vuiou BaDbd(Lin rnd A.a 1988; Nd!rlj|!.!d R ngrsy, l99O; w|!g sxl PlL l9l: Charb.dd al. 2mf,i Mvts .t at'2w 6d 2@5i .nd Mci-aa a 4r., 200s), *iitr M.tldiliw g.c !.d b6 i.pon d by R.tbe ed KbD (t93), w&s .rd pe O99D,N.duj&.!d rdSrt.sy (1992), Mry.rd Cr.o 094), Rdd a at 09'99) &d

'Ih. .&cl of paiob ldrSth o Lrf rhry., L.f oddni@ .!d @.!.q!6rly ir3

EL iD l!. i.ralcprid of &|[ r.didid! i! of gtql6 iupql|e. in bEdin8. colton

lL|!t with gFd6 photo.Fthsic -riviri6, ThG inponlM of paioL tourh |'F@4a gElr6

'r gddns, wni.h hlc rlF pot4ri.l io pred@ bollr .t vdy ely ro<ld.

11 ha b..o obsdd U.r b!9.' pctiolc Lntth (.E l6f!ir. i. bigSpr) f&ilir.ta skndidi@ to FEtrL to th. lnimM, &xl pmvid. Fdrctio! io to[! ton p.!o du. ro

.SrPgr.l

daxnd@ p.netun@ of iroc.ticid$ sthy€d, dd ir @nt&t ndla p.lioL ldetlt myE$dt in rh. dcv.lopm@r of d€@ loliagp cdtirg ir boll rctlenjrg at @li6 !od6 For

pruric.l brccding anotion is lo b. paid io p€tiole lcngltr {hilc nal.jng d.lation of

dEinbL pl&ll itr *gr.ga,tirg populatioN. PBioc in&matio! oD llc g@.tica oa

p.tiol. ldglh i! pldl plodwtion d6 ml .xi* ir lhc liteFturc, &d lhcFforc rh.

inhdildce of thb chd&lq h4 bM qdin.d hr.Exisro* of di'Idar sE is of 6tion laf cul viru (atou €t .r, 198) in

Prtisid sugSc.ts thc nad fd thoFugh studid of gflelic mdhekm co.trclling

qfuD, b.foE d€veloping a bE.dinS preglr@€ aimcd at ddclopirg 6itdt tulerid

Krowl€dge of the gdctic b6is ed hdit bility of Gislan e lo CLCUD is .satial for

fie developndt of Bklrnt cultiv.6. Ths, thc p66t 3tudy wd d.signed to dehincrhc iyD. of aac &tion c@lrolli!8 ln. inhdnan@ of Bistan c io crcuD thmush

gddctid ncm dultsis uing disrue ding sal6, 0-6, d€velopcd bv AltrLr e, dr

(200D and Arhb. atd Khd (2ooD- rtu Mh objdtiv. of 6. Ptt$t nudv ws ro

*min lh. Spnetic nehalint cortnlliig tne CLCUD Gilluce in 66t p'o,,?wtuD L, $ing rh. pldt naldisl dcv€lop.d by msing two cultiv$ drfding wid.lv

for $.i. Fspo's to CrcuD. D.!. or l[. ttgoM of th! pornis, Fr, F,. dd belcF$

sddatios lo CrcuD w@ ddtzcd foUowjns gqedtion ncs daltsid (M.ther sd

,ink, 1982), In ppio6 studiB, thj5 bion ric.l m.lhod had b.a us.d to ltudv lhc

inhaitarce of r6isrecc of gny l4f srol di.@ in con (Co.Ls, 1996; Coal6 dd

whit , 198; ed Th@pe! a al., 198?) .!d b&teri.l blight ir 6lton (wallrce &d El-

Zik l99O), od ths th. @. bionctiic nclhod a 6.d hc !o stldv th. i ditr@ or

CLCuD Gilt nc€ in @lton. ft. infomation repdled hcrein mav b. Neful to b@dd

d*loping A@.tic cbraE ir Cdtpirt ,t6,r!t L agaitBl Cl-cuD, and in

cxploiling vdi.non in donomic plul ch@i@ to d4.loP Pomhine vEi€ti6 wi$

dir.e eist nc. dd good fb€r !!ils u a th. locrl dvilo@@td @ndiioot

REvIEw oF LnERATURE

Itre e vdios dis€as, whicb .ftct th. prdetion Fol6ti.l of grewing

llers in the fi€ldr, Dis66 thai e coBida.d lo b. of litd. inPortdce today may b. in

@idmic fod ionotu*, s illut'at d by CLcuD it P.kitt n, whch ail6t d tne

n rionl .@ony du&g th. 90's. cltuly thc cono bG€dft div.rt d ln.ir addtion io

study rh. lw dis.e in dclail, and dcv.lop rcisidr m.r€ ial of 660?i!t ft]{,tuz L

,.1. conc.Dt of dbse ir pl.nt3

A plnnr is ct?ted halthy d l|lmrt th@ it is sble to csrv out its phFioloSrcal

tuetioM lo tn€ b61 ot it! gadic pohtid Gonidav, 1989) h ! h.alihv Pler' rhe

hdidm.lrc elc divide sd dif,oEdEle &@ding b the t!qui66l! of PLntt for

grevth dd dqelopddt in ord4 10 c&ryin8 oul ttu Bstial Phltiologic.l 4liviti6.

So-irliz.d ceus rberb var.r &d nu.i4t! 6om lhc soil &d prcd@ s*d ln disa!'d

Dldt th. alility of th. pl&l to cmy out lhcc 6qtial fr$ctiort L intcdptcd tv cii!4

r plrbog.ruc nicrc{reanim or d adv6e ovirom4td f@tor (Holli&v' 1989) ln $e

fiRr ins1llce thc .fnidio. ir lealiz.d 10 o. or a fcw cels, and lla not visbl'' bui al

llrer st 86 thc r.a.iid b@om6 norc {id4@d, sd afiftt d Plant patti d.vel@

visible cbsgEs. Th. visibt. chae€. e |pgnizld s tt stDprotu ot lh. dis€s Thu

tll. disdcd plet cd be dcfocd s th€ outcoEe of invisible tnd vhibls FsporoB oI

Dldt ell! to a p.rhogerc nicl€rgrli$ or ihe 6vircmdl5 th!. cllt h adv@

chdg6 in tu&tion of ile ple! dd nay lead to pdiitl inplim.d or dealh of ln' pl6t

or ir. pais (c@q, 196?). Th. titd of @lts ed tis6 tld b@m. disa!'d det@ine

lhe tt?c of phtliological tumtion lo bc dturoptcd, for €xmpl.' 'osin€$

in

phorosyDtheis is ctu.ld by inJetio! of6. fotagc' s n4FB in laf 5?ot5' blightE

rulg nildq& ndai6 eic. In $bc disd6 like creM galt. infdi€d cclls rcspond ro

dvngmdls in diffffil way i... ht!.nrephv d h}!!+tssi4 whi'h tsults in 6€

fotuatio ortuioffi d abnomal oryru (no6.fall ed Cowling' l9?7 - | 980)

q!P!q I

2.2. Pldt vlrurcs $ prllogellc .9.!rr ud ateir n.chrlbn of hv$ionBedo (l9A) d.6n€d viM 4 sutsmimopic inictivc driti6 lhc oultiply

only intra-cellulely ed poidtially padDgoic, On O. olhd hdd, Hation (1964)

d.s.riH viru.s 6 'bils of infotiou. hqldi9 in s.@h of a chrcm@mq 'Ilt* viruls

clny eEnedc infomation d@d.d in then n@bic &id which typicaly sp@ili6 re dDoe prct€ins, T@lalion of ile g@ne or F&sriPiion dd replicalion t kd pl&.

Mdin the hct @ll &d ls sm. oa lh. hosl's bi@hdic.l "nchi.dy'. vi'ls do rcl

capru€ or sbr. ie dd!} ed e rot turctionslly 4tiv. ouisidc th€ir host. They @

i!@foE ps6ite or plhog€ns bot e tur uu.Iy E88d.d 6 gauin micfu4disAnong all paihoAaic orgldsL, Pl&t vills @ th. n6t pbbl.6a[c s they ceedisrrdsine dis@s of eomnic importec. io crept (Agios, 1997).lBmlly, it!ft e72 difimt dcfined go.6 ofpl&l vi|le, 6nptisi!8 ovd 500 s?ei6 inletiry I widc

dry of plsts. Pl&t viruss pos s.nomoue rdge of dift@t pdiicle noDhologi*,

hon sFi6, vato6, gdonc desianoB .nd cxpBion stlregl6 (BN, 199)

On e il b€cue clcd tFt viM6 @d ofi€r nic@Igajsffi e th. ca!!s n|hd

thd th. 6qlls of ptet di..4, €dort3 b.gd ro 3tudv thc mehdiss bv wnich

Dic@reais! cue disdc, Msy palhoaaic toxitu w@ sub!.q@llv dctcled ard

id6ti6ed, a w.r. s.vdl distin tiv. o4h&k6 of etion bv loxiN, 4h .ff4lirg

sD€iEc sit6 on mitochodri4 chloopLds pLsa mcEbffi, spocifc @vm6, or

sD.cific ells sEh s glaid @lh (Asnos, 199?). Ir etldition, Asrios (1997) cmied oul

sad ddaiLd bi@hmicd sludi* io elucidrr€ l[e neheisB bv wnich lotiB aflcl

d kill pl&r ccus, or by wbich 4lls of si6|'nl gaotvp€s .!oid or iMtivata illsc

toxindpatiospnr. M@h.Di@ or induclion of crcM Cl dreac of mev pldts wa

nudi.d b dctail i! l.r. l97os dd 198G. fte studie showcd thar b&t6m(Astobacletiuk t@ela.i^) iitloducd a sPeilic pan or DNA (I-DNA) of ii3 plMid

(li-pleid) ido plad 4lls. Thi. DNA *gnElt beon6 inl.gFrcd ioto lh. Planr @ll

dd is treeribe<t by thc pldl ..U. The T-DNA cont itt wcr.l ga.s, orc of which

co<lcs fd olet hodon€ inilol. &edc &id (LM) 4d @lhd fo. { cr$kinin. O!

qpBio4 th* go6 pFd@ lb. plml gDw$ r.eulrros which 6ult in u@ntollcd

cnldg€ndt sd diyision of c.lls. D€Pdding on lhe clalive con otndon of llt€ two

l0

gowrh EgrLlo6, inf@rion Bults in thc prdund of mryrliz.d dls (tuno6),p&tially org&izcd ratomt' orh.iry mIg (Co!o, 192),

Frem 1950-1980. @dDreh€civ€ dudica w€rc cei.d out on th€ .ff4l of di@.inf€lion on th. re+inlion of host €I! and on tlre posibl. reI. of alt€red cspir.rion in

plut defdcie (Cau&4 19,16 ard Cdickhol, 1963). Sinil&I, iNBde.doB aftmad. aboui ti. pldt @ym.r tnai bsonc &iiv.re4 g?.s ud uomts ofhcbbolitG,

phoolic conpounds ed phcrcl{xidiing cnzyn* eMulaled following inf*tion.

Th* stlrdie pbvided r gdt dr.l of infomation abolt bidhdical @tioB tnar Gtri8aald in pl.rr ells lnq i erioA but djd mr mtirly dplain 0!. nchri@ of

delace ir plels tgrimt th. parhogs.

sincc 19703, *vdl studid hrrc b.d orduct€d to €valute ihc m.tabolc

chhs6 .$ei.t€d witL th. hypdssitiv. 6?oe (HR), r fom of nlid pre8mn d

hosl @ll dqth i! a Fgion mud lha Fi.t of ta$ocBn mEy (Hmond ed Jores,

1996; R'€ls .r zl, r996i vr et al., 1998; Hcath, 2000). ti hyp6euitivc eaocqnmft6 cnzyme knoM s pldt pathos.nBierelal€d (PR) pmteiN d. &tivated.

'ft@ ptlduc.d lR lmtciE ind@ lhe stalhdis of elnyldc, intDdetion of ond.tiE

flzynd .nd preLiB, etimi@bid ffipouds such 6 phttoslqis (Croictlha'L,

1963), dd sme @yn6 that ait&k ed dislve @mponcot6 of tbe cell wcll of lhe

parhosa or @ preteins. irhibiio. tLt n urnlid slein GE De of lh. PailDgEn

(Cdnr4 l9t6), ftc pEtic.l iDpodec of th* Polcis i! aa€ri..lly hodifed

pldts i! thst, upon bferio., they will podu@ suE€iat douls ofPR pDlciN 10

otu141 lhd fron disees,

l3.Vitu blddo! ud nplkrt'b! L t6l e|lPl&i vi!l@ 6t .1h. cells oDly tnough wouds Dadc chmically d by veio6

or by deposition inio m ovul. by idded pollo eFiE (MinI, 1993i lohsFn er ar,

194} E$ 0 9?D dd Eofd d al (t97) 3Mi.d lhe Epli..tio! pmc* ofbet *6lmy.[ov viru, . srll i$m.ttic viM TIEy ruSg6ted 0u! viM prniclG d fi$r s@ in

marw sid. d44t5. L.td the viru D.niclca appd in prcrchlm! cclls nqi ioplasmdeMAr. .omsting lh. pamchFa cell5 win tn. mat@ siev. .l@mlq

$@npsi.d by thc dd.lopmar ofv6i.la @oLining nEl.ic &idt Th. vdiclB @

u

q.rp!9.'...-... . ..............- @!tr9tgt9n!s!-

tulos.d i! cndoplMic rdicdlo (ER) cinc.n!.. Th. v6i.l6 ed thc nwt.r' cnv.topc

re lrt .hcd pith 4h othd dd vic pdricl6 app@ itr rlc treLur lai ro tbe

nEl6lu. A! th. tihc of viru iltc@6 p.nicla r[@ s.d&. thrcughour rhc ruql4 ed

Ihc au0r€rdc nc$od of Eplication of plmt sinetesttutt€d DNA ($DNA)

wr bs noi '€l

bca delmin€d. nr.rc is sme wid€nce. howevd. rb.r th. ssDNA

Eplicltd by fomins a relling cirl€ ih.i poduc6 s nrrtinaic c) srud, which tud a !dpl.t! for thc productio! of nuttindic (+) srad! lhal tho clovod ro pDdMuit l@si[ (+) st&ds (Mal!ds, l99t). And tbc prcdEdo of w vi6] n@l.ic eid$ne is Fr5la!c4 iftl@iry ln. hosl c.ll to Foduc. thc poroin noteul6. T1F prcr€jns

fmcd e for .xcluiee E by lh. vi@ d a .od or in ofiq nnctions, \tto nd vituneleic &id dd virus pmlein subuits hav. b6 6yn1h6ized, rhe neleic eid orgejz6th€ pFi.in sbuirs eud it, ed th! two N Gmbled log€thd to fom rhc chpl.Lviru pcniclc, ihc vinor h $DNA vilrs lynthcsis ofnwl€ic &id ed pFt.io, s well

6lhcir .lmbly into viriou, tak6 pl&c in thc nudels ton whicn rbc viG omicldre th@ Eles.d irro d. cytoplatu (Hull, 1989). ftc 6rs i.lacr vino6 apFe h planr

@lb .ppoxin.r.ly l0 bo6 lna i@ut.!on. n! vi@ panictB may di.l si!8ty o ingouls dd nay fom doDlou or cry$stlin ifttuion bodis Mrhin th. c.[ cyroplsm

or nwl.a (Chrirtie &d Edwado. l9?n.

2.4 G.nlnl Ylrus - Mode ot TmtDkdo!, IDpr.a otr yt€ld rnd hbcrtr.leStldlo on Glob.l As&

All ihc BcgoDovi@ @ resincd by wlitelly veroE in a pdktat,ci@ladrc nll|rM i.e, ore l[c v*ror fc6dj on e inlet d hosr ptdr ud &qdr6 th€

viru, trdsisior of th€ vitu tu rtd aN qifiir hoN, ad m.y @ntiruc for rhe tifc

sre of th. vetor (Coh6 &d liaed, t964). Howevn Coh@ atrd Hlrpe (1964)

Eport d ddlind in trmnision .mdmy in tomato ycllow laf cul vjN! Oan C\4,whilst Coha md Ardsnus (1994). Potsro! ud And@n (tF?), Hirje .r .t, (2oot) ddPalubo €r al., (200t) r@orl€d @nirol n@uH ir i ared Fgiotu by hptsizinev*lor @niDl mdnly hy p6tici.l6 or phylic.l bdi6, Chmical mntDt ncibodg n!v.ba oDly p.nially cfretve, sie whii.fly populadoG m I@h vdy hi8h Dmbq,

t2

.91fP.99! 1. ...................-... R.d4orljir.nt!&

lc.ding to intersiv€ p$ricftt uc (soncrimcs twic. dail, iD anemprs ro €timinare rhc

v41or b.foE it lresnits tbe vinq ed rho ber w.y ro rcdu.. goiniviru dd.gc i3 by

brccdiDg mps 6i!ia!r d bltur lo ihc viru (Cob6 ed AnriSDur, 1994., pi@ a at.

196; ad Moal€n 2001)- Thc irhdirea of 6idm@ to begmoviM nad hd!rbdi.d for ndy ,€rs, ald evd.l loltG ofsbr.s! p@ shdm to b. nulti8aic ircie. A praldt prcbld in hr..dirg for si5lre. b cdini vitrs h.d hcn rtEl&l of @Bisr@y in vinl @btuLlur!, diflc.lnr vitrB hav6 iI6 s.fu nM., udtm6 of vills t€€p or cl@ginr; &orynr th.r l@k like lhey inply slrriB &ru.lty3ignify vcry diffeMr villls. (Fdqud ,r ar, 20m). T[€ I!@1 findings slEw rhal

eonbimrid betwed !p€.i6 of gcninivirus hrpp@ rcl.rivety ftequotly ad !d&!o t!€ already conplcx situalior (Padidm ./ at, l99q ed Fequd d at, 2000). rthhos c.uled .ontusion for brccdcf ed Gehasr maling .omuicarion ditticulr,

slowirg prcger and cauing i@rel srDptios,

Yi.ld loss due 10 thc tonab ,rllow laf cul vitu CIYLC\D on@ t@h I OO%

(Pi@ .t al, 1996). fte tt'e TYTCV hd ben eim 10 t ld@gmB c6pld ofbc8@ovi@ (Mdio@ ald Caniolo,20@). TYrcV h.d b6 tmm fo.loE in rtcMiddlc Est No.rh dd Catrrl Afrio snd Sourhqd Alia llte dis.4 @.li4cd

Aradilg to $ulnd EurcD€ *b@ s6 ourbial! hNe b@ E?o.r€d (Ca$d( rqlt!rm! l9?; aDd Natlla and Mawcu, 198), IE erorte to blEd for ryLCV-Gistet qrltiw have t€d thc da.lopn.nr of m .pprepriale viM iljMbdo! ddscEoing netlDd to as$ lYLV rcdri@c (Lspidor ,/ ar. 194.

In wlis snrdi$, inlolvirg l0-&,5{ld s..dli4s axpoled io tdge nmber ofviruliffrus whit di6 wft reponcd io be tOO% .mdol in ttusfdirg ft€ virulL^piaot et at., l9O. PregB i! bEdinS for TyrcV Fskta!@ had bd 6low,

prindily b6.@ of the oDlld g@ric nAlur of jniqira&e of gd6 ontolilg6i.rssq Th. e of . Eli.lL D€thod fd mina gmplM against rhe viru ig

d.pad€dt on th. .vailalility of vidlifftu *ni.0i6 (bpidor et at.7991, ttvidlv.ky €' ar., l9E). Mdy leeh wo*d hld i?o €d dlt! oD tne lMbd of g6*@mollins rhc 6isr4€ ro TYLcv for q.dplq Fri€dmd €r ar o99E) foud rtullhE bt€tutiry sa6 wft 6!oEibl., whilst vid.*ky ed Czm.k (1998) @oncdtwo b ilF &ldilivc l!.siw gm4 io bc iovotv.d, nF sludy otviitavsry er al. 0 998)

l3

.{&P-!9r-?

ha shom the prcmcc of one nqjor goq A nsjo. 8e!e tcmed 'TY-l " ed at l@l two

morc modifid g€a6 ddlrkqd by Zmir .r al (1994), .nd stuon er.l (2000) fo6dtlut two gmq with rcn-dlelic iniaa.lion @nrrolled .olisbnc€.

Begodoyitue€ in.ssw ce cN* yicld los6 up to 95 % (rrggi, 1999) along

with its murl lo.es BtiDat d 3t 30 % (F.€cit er al, 1996), An epiddic in

Marlaeee in thc 1930r dmorl .lininltGd rh..ultiv.rion of cssv., .!d l.d !o t!.8En4.tion ed d€v€lopndt ofBilrlar vsidiq fi@ inrd-s?4i6c ctuFs (coN .r zl.,

197). the incidde of tnc di!.e& @iftid.d with th. whitcly popul.tio@ ed Bi.fall

.rrl. in the,€d (L.sg rld Thct! 2000). No adcqld.ld.l of Gblancc ro ClsUMolaic Di!4. (CMD) w foud in th..ultivat d .rs!a s?di.\ Manihot adlenta.

R6ielaF. wa lals intDga.d fron olld cbt d ,lart or spoi6 by inle@€.ifc

@s 01.!r a at, 1980). R6i.lac. non uE intcnp€iac @s i! c.gu is Efoded

to bc lE6irc, polygdic .!d lddiriv. wirh . hdihbility of alout 6 (H.bn.,a1., |980

ddl989)

25 Vi.d di6. oa@lr.r

25.1. Men.dro of 3pMd ol cotto. l6f cddpb vlru.Cotlon laf cMrlc viru (CtC l, t biplrlilc bc8omoviu (Nadm ?,

"r.199, we foud 1o be ei.,r.d wilh rh. crulDlc d!!.e i! @t!on (BmM .nd N.l$Al9E4; ed Bdddo! &d M{$d, 2000), Conon cMpl. di3ca @idy .f,.c15 cotto! i!lh. euthm USA 6d ..!s rcl.tie.ly nild .ynplotu 6i.* it uu.lly app€a late

duing the otLtr gbwira d.r!on. lrs Eultiry fron .otlon laf mbple viN(Cr-&\,) i.fatiotr h.v. bq t!!on d io dgc tw 2l to E6.0 % litc cotro laf culviu di!@, d@odi!8 on thc age of pldE (AIlo.r 4a, 1960; BrcM.l ai., 1987; ed

CMple vtus of @tton hld bcar r.dlniticd dpdihdt llybydaso /ard.i to

ndtu6 pbDt s!tui.s witti! ll!. M.lveq. ed F.!.c.ao fmili€s (BrcM dd N€ls.n,

1987) lik cotion l.d cul viM dtu@c, n ! my .xplu why ht€ding conon for

Gi.lance 10 CLCrv ha bcn prcgssing ntha rlowly, Ir ha<l ben sugg€sred lhat the

high CLCrv sbidce in C.dix, @tion vdidy fiom El S.lvador ie controlLd hy two

t4

.qrPir:

dotninsl od supplmdbry sfres (Wlson dd Brcr4 l99l). R6isb.r lines, howacr.yicldcd rebtiv€ly low nd complicating rh. d.v.lopn6t of resi.ter pldi natciiob,

R@dtly, mor€ cfforrs w@ nlde 10 6nd CrcrV rcaki.n@ in upland @iton. Dle !o li.kof s .lt hativc ir@ulation srslem (N.twicl er al, 2000), rhe jts€ehd had b Ely @

fi.ld infelioA which hoqe6, hilda.d ih. preg6 in iddtirying Bistant gmdic

2.51. lDp.ca oa CoLon L.l..rl vinr (CLC!D) on plri prDdlcdor o! gtob.t

\vhicfly-rdmitd Cffini viru-ieit.d dis.lg clhs in rd!.ed grc*1h .Dd

siSnili.mt Fductior in yi.ld .!d qulity ofonon- Virus inrelion a1 €dly s|.g6 ot pldrth.lopmor r$ults in lihl€ ro rc prcduciion id nG1 widely groM @non crlivs,Coto lcaf cul viru! ! di!@ having $a poldtial lo cFale disslc. 10 otionpoductiotr w6 661report€d on G8srpiud borb!.le8e iNi,gena Gaquleon, t9l2),.t66 $d M@n (1920 ed B.ilay 0934) r.todcd l!!t CLuD had b@ olc of itrc

no$ sioE dis.as in T@ic.rd Suda GtetiEly, ud ciqrEd bajor d@r6 io@tt@ obdElion in the two 6uEi6,

In USS& laf tul dis.e h!6 bcn lsDM to ca@ oajor redlclion in nmba orbolb of up ro 55 % dd 6 bign a 72,9 % whd inldarior @urEd *ty io ihc s.en(Mo!kov.t, 1944). Mlh avd8. i.ld los of t6.2 10 t5 %. h additi@ to rh. i.ldGrluction th. die6e als afi@t€d rb. qualiiy of cofion fibre rEd@ing ldglh by about

7,5% dd slr.nglh by 3.3 to 16.6 % dcpoding on ttt€ vdieti6, rrwin (1927) at& foedlhd qtcnt of danag€ depad€d on ihc a8c of thc pldt at time of i.f6tion. lf th. pleis

inf6l.d al vdy @ly )oug s@dline n g., rh. radiy of CLCID Bult d ir sruniing ofplsr gretl! ed ultini-ly dw.l@ Inb or m boll fomatior Ieb.n (1929) rqodcd rlo$ of 20 % h yi.ld but eirb @ cficr otr gFdc. Massey (t934) nDrtc s q@ribriwdditr{i@ of dise* ard shovcd Edrcnd in i.ld !p ro 50 % d a@ur of CLuD.Atdws (1936) fourd l6rt thi5 CLC\D Educ.d *ei8ht ofopa€d bol|s dd .wbd of

8@o bolk by /{1.67c Dmb6 of bud! ud flowd by 4.3 o/o. Tm 0 95?) st t d that l..fcurl di*. in Gei6 a!!d Sudd cNlcd l0 % io 40 % ield los6, Esulring 15000

l5

.S!.P!+:

los$s in conm bal.c. Lanbcd (199) r€portcd I toss of 20 % itr i.td, by omparing

hcdthy ard inl€ted pleb, but rc efrer on 8i.d..

Obserioc by vrioB l*thh idic.re rh.r ono p|das qft Dod

N.9ti6l. d 2, l, .!td 6-wch old st g6 (El-Nou dnd Abu Sglih 1966; El-Nou. l 7

ed A}ntlr a al., 2004). ?inl (1975) Epon d $at tnc ldf cul mos.ic viu @mpld inth. Aticd @lton b.lt ws Elar€d b fou di@s; ldf cul, sall dd t{g. vcin

ihictdjng ed mosic. C.rqil !trd Folli! ( 1983) hrd a @npr.h6iv. diNion d the

t)@prorq c{|ls; nod. of tlilMi$ioo, dan g. €us.d dd otrrrol ballG of the

Ani.e l.d cul, AD.ric& laf duDL, moeiq bluc di!.e &d olha lcsf b[

2JJ. Eblory of rlrdopDdt of oatou L.f .!d vL!. dtt.&(CLCtD) i! p.t!r.!N.G hs ! .i!8lc pdnoga or iler p61 thetqqd cotton .dhe h rh.

@utry, a had lhe cott@ lafcul viru diee (CLCuD), Th. lcsfcul diee of.otorwa liEi obw€d on a fca bdiidual pler! n d Muloa (P!Hsi.r) in 196? (HBainad Ali, 1975), &d conrinD6 to Ddsin on rhc @non cDp. Thc di!.& badepoDiMr in l9?3 whd i|5 plere % rcrd on eid€ly groM vdi.ti6 l4CF rd B-J5?. Di!.e ilciiLrcc on rh6c vdi€tid vs aloui 5% (T&v6, 19?4), Th. lowqincid.ne wa @inly duc 10 .tr.chng conotr pldl tate ir ilE a.3en on rhe uppd ponion

ofthc ph . b 1987, $c i!cid@* is.h.d up io 8oyo in crr.is fi.ftts ard 60 h6cL6 of@!o! w@ ih'g.d i! Mllru disicr (Hleb ard M.trDod, 1988). I! 1989. 2OO

heris wft !f6L4 ed 800 herarB of@tlotr 6.[4s w@ rcponed wilh CLCuD ir1990, Th. ituideE€ iercled slbdleliatty dd cued 10$6 mSins 22.3 %ro 68,5 %

in .flEtcd 6dds in Pwjab Pnvi!.c wi0r d.p.rdilg upon il| ldicly' rjo. of infetjon.!d @viFl||ht r @!ditio6 (A@,rDou!, l99G9r). A@i n8 ro th. r?ons of Oilrud Nou (1969), Kh{ ., ar (198), I<b6 &d Khe (2ooo), Atdt6 et at. l2@3) ,AShm. sd F$hi (2004), th€ idpont p.t|@n for the $Erd of CICID i, hiShly

dcpaddt o t np.rdur., hMidity. v@tor poputation drd th. prevailine vihl lo.d_

F.gctlc ., dL 093) Als rported driB f-ros Esposiblc in ite spmd of rtcltlpi.d whiL0y-tralmincd gtuiviG of dif.r6r mp6 iD di!@ dvilDual'

t6

!n 1991, the di!.e pl&ying a major nlc io h€ cxlmr of hfccrion ov€r 14,000

h*tac w@ aif*ted iD Mullar, Khedd ed V.hei Dislricrs. Duing the t@ 1991,

A€ high€sl otton p@ducdon ws t2,82 niltion b.t* ir P*ble th.r mly be nu€h

h€igbd ifdiw did noi aII€c1, h t92, Ur. di!.r& ws r.poned on norc thD 48,500

hele.& cauine significet Educrio. in prcduciioq tud @onomic lo$ for rhe @dtry.By 1993, th. die6e h.d spErd to ihc .nriE colion b.lr in the Pujab rDvim.. vsyineirt tr3ity dd danage w3 obery.d to rhc @rlon crep on 889,000 h*tjB, dsr€6i4@iton yield b 9.05 nillio. balc!, which wB furthd r.dEqd lo 8.04 milton bd€s ir1994. latbr.tr d-lar€d a distd i! @trotr pbducrion in 1993 (Bb*T, 196), due io

high disee wdity ed widqQFad ofthc di!.l& be.e ofhgh phiteAy poputario.

In Palind! 6to! l.af cul vi$ dis4. caus.d Djor rducrior in .onopmductio! of7.l nillio b.l6 (duing r!. Lsr d.cad.); which c!u!.d ! los ofRr t_2

bilion dolsE to ottion:t @l@y (M.!n@4 199). nE lpiddic or CICID i!Pa&is& i3 o& oftE bcst q.rnpL. of rhc dautic shin ir ibport !q of. prwioBly

uibpoirlt di!!'@ (Md$!e, 1992 at Z.ls d aL, 197I

25J. StDlaomloa, ot L.f.rrl..d cnDplc vlnt dl'.!G tn on rThe tl@ of thc di!a* w 6!n le.d 4 ..lef .ut" by Ki*p.licr (1931) ebo

al$ {t 66ib€d th. sybpiob! of th. cototr l.sf cwl viu. A@rding ro rlc !ldno!, rhe

stDploc of rhe di3€e may v.ry .@dir8 !o v8i.ry. s4aly hf@red ptols myslow s?irdly rwilt d laf p€lioL.r fruidng btmhe e4 b . t6s qtdt, l[. mirs|m, tends b gbw t ll wittr .longl.d iordodcs in Oar9"r'@ barbad.$e W. G8,1951). ft. dis.& is ch.dan.d by cirhcr upwad or doMMrd .uli.g ofl[€ lav6(B.i&lon dd M.rk!.r\ 2000). v.inr of thc lqv6 bcomc rhi.k€n€d whicn N noEpDrcucd @ rhc ud6idc,

Two R!6 of v.in lhick€nins e comonly *o, shall vcin rhictsing (syDed Dain voin thickdins (MVT). Sndl v.in rhjckoin8 is m@ comon ude li.td@ndiiios dd is ch&&taiz.d by m.ll gtq bcad-likc thickdinS or th€

'ougqlavca, The* iftgul& thickdings gradudly cxLnd ed codc$e to fom ! ootrtinDus

r€liolation of ihc m.ll v.ine. Main v.in ihickding is ch!rut .ikd by ihe eI@lhidding oftnc dkt l dd! of lhc lstd v.ind of youg lev6. ftc thickaine fi6t

t1

.!+|P.!9.'|

4pcN ree th.lofnargi. and rhen exlcrds in{,rd ro fonn n netwo* of da* seen

0ict@.d @ir vein (wa&na, l98l). Ir dbd. but ml in&.q!dt cascq fodatioa ofth. cup i[+.d' o.l€I lditu orSrc*d cdl.d "dtion" rpp@s on thc urddide ofthe lq.t En liont hlve howda ba@ obwcd €v@ on lavs ot plrnt with only nildbad-like vcin thict@ing (MdD@4 1999). Ac@rdins io MoErafa 0961), clniDrtDplo@ qp.sd on th. rop l@16 30 dlF lna plartinS rnd i dsi(y r..ch€d upro

100% within 15 d.rs, It hld h.a studi.d rh.t rhc aymproms in @tioa Budly @pcdwitrin 2-3 wek of in@ll.tion by B. taboci (Sindn et at., 1997 j ed Alltd .r al, 2002

a), ald @ pdMily ch@rdiz.d by. d..p &Mw{d @pping of ihc }ouleest tov6.AI ile v&i.ri6 slow dw.rfDg .|lat eha ,ttr.k€d by virus" moc e in rhc d*rfvdi.ti6 lik S-12 ard CIM-70. InlerFdsl distme b r.duced and Ih..lfed.d dehbome snbt d in @ly infetioD wilh .rlv@. efd on Auidns. Ib. cd@tio j!!ub6 of bolb rd boll wciA 6rnr in lG oa pbtrt yicld. h cxrm. cc, tbe pldts

ftc atmpiom of cotton lqf cMplc viru w@ tiEl iine rcpodcd in Califomia

(Dictm .r dl. 1954), .d t fs le8 brd in Ari@ (AIs e, a/., 1960). A[!t ofcoio lal cmd. viru (Ctlr\|) is ch@rdiz€d by floEl dilrortio4 hrp.iuophy ofint rveind ti$uc @ltins in dohwad culing of lov.& dd foliu nosaic ecompeied

by v€in cldirg .rd n€qu@t vcin disrdtion (BoM e/ al., 1984.

2,5.5. Vdto6 aor dE rror! slo! ofCLClD rd.lt F.r. bcttColging (1930) d.noronztcd tn sympiom of di!@ ddelopmdt wirhin n-24

dlts vho wl r.lly wa u5.d a . rero., ald ir additio!, hG rlso r€toned tral j4id ws@r r v*tor of lh€ dis€@ in Sourhd Ni8di.. Ysin &'d El-Nou (1970) ffitullylrumitled this disNe by a single wlicny, Hoplis (1936) .tenonsir.Ld succ.stul

llllmisi@ of iI€ le3f cul diee tom wild speis of Si& to @tton by bud\4irs ddgbnbg I( es ob6@.d lhrr the viru @uld b. rr@incd toD infercd ro helthy

Phnts tbtuuih gtsni!8 .!d lhsug! wnnGfly, ed lEeision ilrcueh gnning pmv€d

suc@stul (Miu4 1992). Mia (192) d& r.poned tnd conor laf cul viru wsn iild n-hdic.lly ta@ind nor cui.d in sil or rhFush se€d. S@ r al., (2000)

nrlld rFon d lh. dd6ti6 hdhods for rh. prEe@ d .!6@ of 6G viru!.s, dd

l8

foud th3t CLiD w6 not ttursnisibL n chadc.lly dd d infslioB clo@ k s fdnot .vaibrrrc. Accordins ro Brcu (1996) th. Bmisi4 toba.i (Gsn.) c@pld,

onFting Iob.@, @tloD, lwer por o q.'hirenr, ad B, ^tAeatdolli,

or th. B biotyp. of

,airtz rara.', re lhc otrly looh whiGlly qio6 of sbgrelp m 8@i!ivitlB, Arar,.t is €8lily calat'ls of stallbhirg .xiian€ pofltlarion lry.ls" pa.ticuL.ly in cbF

8roq udq irigat d, eid conditioft in both li€ld dd gMnhose syslebs. In.ddirion,

ihi! whitcfly h6 the poldtial ro coloni& | wid. Egc of djcoq4.domq spaid,priEily v.gltabl. dd 6br. 3pei6 wni.h of grd iryon4. b mrllwi<lc agriculturBl

Fodu.lior R€@r studid indiclr€d l[al thd w€E nMdu popuLtios oa rdit,a,ara.i th.t vary in ibeir capeiry to d€v.lop high population &ns i6, in hosr Mgq ddthe cflicey with which th.y tr@it g.miniviNs (Bcdfo.d e, dl, 1994; BDM ddBird ,1994 BreM er al., 1995!, b; &d Abl& d at , 2002b).

Conor lelfcul di@. k tmniu.d by fe€dila whit fl, ,, ,ara..i witnin 6.5

ho6 md a 3inel€ f€malc, cdying vior cs j.f@t mdy plet!. wlirelly ie tloM to

swivc on 4 Doy a 53 h6t planr Aei6, dd tresir5 about 23 pl&t dis66, Atglobal ld.l, wbire fly irf6ts 6@ difdal ptdt +ei6.

Alt tuite ho6:r plqc play a sigrifcdt rcle in dE s?€!d of CLCID io coiion

cop, Thcy povide inoculum 10 the veror foi trrlmitiing vitur to @rion. A runbq ofpldls havc ba Mogniz.d & alt@te h6ts of Cll\rD. HBlin (1932) Epod.d lhat

lhc l6f cul dtu es pr.$qlr on gada zinit in Nodnm lldia &d it {s iddrical ,o

thrt @ord.d on @ttoo in S!dd. CowLtd (1934) Gpon.d ll! t thc dls.e wB HdilytrMisrible to @tton t n tirittu etelentss 8d Eibis$ can art6. Mckovdt

( r9a0) in USSR reponcd tlibbd aMUn6 Nd S'IMM d4l@ndra s altcmatc hons

of @tto ldf cul viru. Te (1955) @td.d ir' ptErc o. Dolitu lablab, Ph6@lB

trilobut tA g@ e@ @, ntneo\, lrd in 1957 be again stat d rhd Hibitu 6cuL,tus,

H. nb.loifa $d H. .a,,arin6 sfled a the pdmary s@.s of in@ulm of dris

dis@c to cotror Hs.in &d Mrhsood (1988) found thai in ?akiria! thcrc e nore

ilm 40 ilE alt€mde hcls ol CltxD, allhoud rh. dact iddtifclfid of Oc vitu or th€

pldr spci6 w$ d.lu.&d.

l9

.qr.p!!Jr........- ......... .. ..-...............I9vi!]1erll!q..9."

Mitu a, at (199"1) in @llrborarion with vircloSy !.ctid ofAyub AEicultrtrc

Res@h hstitute, F.tual.bad (Pakistar) u.d ELISA &rr b d.lulre tllenaic hosls of

cLuD. ft. autbor foud tl,l CL{UD rneked a wid. vdi.ty of allalarivc holrs

h€loDging ra sad tuilia of dioa s wll a ltll)|rDt Of tb. 6&r6t d pler 3pei6

$|a.d uing EUSA, 6l 8rv. pcitiv. ButB of lt€ Pc4. of Cl,cuD, h 195,

Malit .r ol ur.d polF@. chrin rc&tion (PCR) dd DNA pbbr r€hrology to idmtify

dErnar. hocrs. He r€tod.d tld I'iri'.4 6@lar4 (orra), r, r66t@tr (chim o@),

1' @.ebh6 ('surlrn'), t{ tilio!@ l't E i'\ fuqatw t\d &ida w@ .llanale

h6ts of Ct.cuD, wbilc chilli6, ioturo, bbuo od zimi. w@ mt.

Hu$n e, ar. (2003) $inc suihn-blotiing t€chniquc confmed th. pcqce ofCLCUD DNA p in chjlli plpF pi,r$. Synplom inrclvcd lsf .ulirg (l,C) and v.irtnicloi!8 0/D fd tbi! CtalD.

2.5!6, lt v.lopmtrt of CLCqD Bbttrt v.rLrl6 o! glob.l bulr md ir P*hbnAnd l!€ oMt of @t1or lal cul viru disc much cfrorts w@ Ddc by

tEeEbc- to brc€d for CLC!D Bi5ilrt vaiclid in Sudd .rd Nisaia E&Flis SaIsl.d @t!oB sw g6.6lly cui\dqld se.?tible; hoyad, Gisldr pl6r nr.ri.lm &lcd.d tom the Egnrih vddy, Sak lin 1920. Ilmb.n {1929) iddtificd CrcuD

Gistalt pldr mareri.l tlh ihe fi€ld of S.k l in 1924 lad daelop.d two Bislsntv&icli6 of 6tron, X1530 .rd X1730- Th. .ftctivcN of O. *leli$ ha b€EairLi!.d fd Dcrbe 30le$ (ci[' !d Nou, 1969)-

la Sude, elsiion of individul pldts non sgrc8aiing pQ'rlalio@ undd i!€high CICUD incoloh lord w3 n.d. in !.eh of highly CLCUD qisr.ai lifts;howdd, @ of the slort @irgs of rbd clerioG B itrfaior 6ba qu.liiy. Siddiq

(t968) md. cM bct*a salel (a [igh 6ba qulity vddy) ed teb.n (t tow

fiba qu.tity Eiety wilh lcaf curl Esistd$.)

h Pllis1!4 thc 6Bt rt p wa 10 id61ii/tolefuteirL cr tir@ tm qidtine

e@ of Aa*tpi6 hircuttn L PreDisirg 6Mi.l cultivN w@ pldtcd in .6tr{)m ro bc high risk, No culrivG dhibit d @bptetc siuimq hoswa' vri€ti6dill@d in rold{@ to CLuD conplq. Thc diFe saairy wE high on two vcrictid,

20

qlrP.!:r.f

$12 8d cIM-?o. ln lrldirid! . vei.ty tmc R.hbei iom $e Sirdh Provilce s$ slso

foud lo be hidly $*eptibL. h c@tret, v&iai€s, SLfi-41, CIM-240, MNH-14?, Blt-

36, FH-8?, MNH-93 ed CM-109 wft ldsr llteled by CICuD (A,ti et o1., 1992).

B6id6 D.tirg $l6tion in qistilg culriv.t!, .ffdt5 9@ b|d. ro bGGd wsirdtu wnn Girtee ro CrclD, Two doric vsi.ti6 CP{ 52 ed LRA-5166 cxhibit€d

Sood rGisltre io CtCuD, ed th*foE w@ ued s pMts in rhc bt{ddiz.rion

Pre8r@ Itc tvo prenisn8 li!6 i,.,1098 .!d I | 00 @ltitrg tn rhc .Gs, 49287! CP-152 i! 1[. cbp sMn 192-91 dldbit.d b.nd bl.fuc€ to CLCuD, Sinildly the

@@ betwd l@.1ed qotic vitu tol@t cdtivs w.re iniriar.d !i Cdtral Cotion

R6@h I!ifn!G, Mult& @ati!ro) cutritr8 in tn dqdopDdr of l9o viro Birr.nrveiai6, CIMn 100 .nd ClM-448. Th6. rwo v{idi6 w@ app6v.d by th. pujab

Se.d Coucil in 1996 for sa.nl culrivari@ b the Pujal, Pslisru.

Addition l b.!.di.g worl Drde ni trc ffi CM-109 x LRA-5I66 ed LRA-5166 x Slz l.d to rh€ dp.lopmdr ofcullivsr, CM-443 ed CM-446 rtDt w@ t!r6appreved in 1998, To sle€d up brcedirg efforrs, th. conon brE€d6 uriliz.d ite avajtabl.

Gisirdt gadic @ul6 of CrclD;.trd d4top.d $tr.l wieri6; MNE 552, CIM-,148, C[v!-482, yH-53, Flt-634, FH-900, FH-9OI &d FH-I0OO, which I.d lo ift@econon poductior in Patjsiln. Alrholgh rhe vdidid €xlibiled Eisr.le aglitrsl

CrcuD bul po$ed sl4 ddsits fd cxdtle, poor @togicat pl$licity, .rbdccpo$e to aviremcnrd .!!s€ ad poor 6bq qudny v.lE (cow 668) whi.hw@ Mjor hurdLs for ft€n adaptlrior lo major cotion gbwing al4, Ii ral ys,wi.ti.q CM{73, NIAB-999, CIM-499, CIM,?07 8d NtAB-ul. .ll p@irgCLCUD 6i.1aec eith impbv.d 6brc qudny tnirs we rteded for rdail cuttivrtion

2.5,7. C.i.d..tlldl6 of otto! t .f clrt vtor .tis.eTh@ e fw Elott! on th€ g@dics of lalcul viru of @rion, Appro&h6 of

brE ding @tlon vaidi6 6isr. lo l€fcul virE &d thc intdiralc. ofctauD hlrtbd r+ort d by wious wo*6 (Ali, l999i R.nmd d al, 2OOl; 8d R.hm& ., ol,2002), A mjo! goc with additiv€ eft@r id relorled ro controll ldf cul rcsirte@(Knigh! 1948). Siddjq (!970) conduct€d g6.tic sludiE of @t0on t.gf cul vitu uing

2l

.g$P!sr? ......... ...... !fytfy9f.-!!q$+P.

two slqplibl€ rnd two rcsilta!1 cotrDn varic1i6, R6ultl, showcd rhal a mjo. dooiMrs6c @EDIed cisl.ft.. HoM, siddiq (l9zq .r$ found nie. (nodi66) soc

AIi (199) lhow.d rhal P, poputarion gav. e qc.ll€nl ril of } I ratio suggcaiinS

ln.l couon IaINI vitue ws cortroued by a ringte ttomi8t gcoc. On the ooE hand,Rsdr'lwa (199) .@on d rhar the dis.e wa @nEotted by rwo donin"rt sad,dod.d by th. n.lc &d fdat. pamr.

h e otlrs ltudy lqhal er ar, (2003) fomd duplicaL dominml @islais io bcinlolved in .ort@Uin8 of CLCuD r€sisLnc, Fv.aling th€ invotv.mor of rwo domiMlduplic.rc ga6 on rh. bsis of bo<tific.riotr of rcdd 9:3:3: I F, ndo lo l5l: I ud r.sr@s *a Dodified b 3: I ririo inslad of l: L Finditrgs ofR.hDs .t a!. l2c{,51@.ddthd 0'rc g.@ w@ involv€d in G ,retun cisr.ne b CLCuD, rwo to. ci3t!r.(x,qo& &d x,aod) dd a suppress of ftsislee (,taddr)_ On th€ oths hed,i.}uj^ et a|. (2001) Eported invotvmdr of two gds with dupticate donindi,domitut i.nrlirory &d dupticat tlBiv. Gtistalis and rh@ go6 witn i.iptiorcdohi.dr cpisr.* h @nrotli4 CE\rD Bj6ree.

Iihairdc. rld holslte sludi.s of disc4c sis&rr g€ns in cltriyat€d @non

show.d that Gisrcrcc to cotror lqfcul viru in c tr&r!4 L. qs coftollcd bv a

singl. donitut ga., wirh no cytopheic .Ifor Graida,2002). R hm.r dl (2OOl)

ob6ad thrr 6isld@ ld.l in die6l fin6 of |ptad @nor w4 .tT61ed by )ts duc

io diil:M6 ir awilsllc iMolu load. How.vd, th6 6dt .!Don! hlye inrtic.t d itEtr.risidce @y have broka do!4 eirce @ilrdr vdi€ries cxhibitcd CrclD sFrDrom!

in th. Vehoi disFicl of Paldsr.r (Maeorcr dt, 2OO3 a; &d Malmood e, al, 2OO3}

Tbc t!3@nm @riddon 4 ar, 20OO; .nd Rehtu er dr., zOOl) filihd r.poned lha lh.cullivs qft ll.llw .d.pr.bility io difaat gre*ing @ndirioG, gwc les i.ld lhquiniotcd su$Etiblc crlri6, A.@rdi!g io Zaou er al (1998), thc qi5t.M ofdift ert straiB of colton lcd Nl vitur in pakisran sugedtr rh. n*d for tnoeughstudi.s of Coetic deheisn coltolling CttuD, befor d.v.toping r bMdinSprclrhmc ained 3l &v.lopins Hirlant nlrsirt. th. avaitabil y of a efficidr E[.!1.d'!a$ s.!qir\g rebniqu., whicb @uld idatify rhc 6isLni dt rid, is inDdranr for

22

.g|F.!r:....-..- R€viaof Ltr.rnlE

good sw* ofplanr inpblaDor for g6c!, .,g, the re.hniq@ srrcss€d by Tripalli mdVma (2003) in id@tincAti@ of sw@t good 6oucs of i€rilhnce to the lomlio leafdl viru, ft. i!0Er rludi.6, rhic! @uld dpt in ihe g6.tic b6is of 6ist4. ofC[,CUD .nd orhd €o@mic rrirx e tuntd Equired

2J.8. MoLc r. rtudtB for conor t.tf curt vlru dtsr*Ia ordd 10 uddtrd th€ nltw dd tbe Elatio$hiD of rhis vitu! wiih oftd

C@ini eills, prnicuhly Ctcrv, N.d6! €f a/. (tB?) !s.d potyb6r5. .hrio@lion (PCR) to upUrying the CrcuD DNA. For lh. dplificario. of DNA r{ of dtwnn.fly 1tllMin.d c€Dini vitu6 of CLCuD A6ifc pdm.B aft u$d. A! . rclult.t*o DNA FraD@rr of $oui 1.2 tb ed 1.5 tb w@ hDlif.d. h thc m. pCR

@d@s t*o DNA t€IMrs of d!. CICrV DNA A @6!M6rs *w .tso dpliti.dft@ t*! n?gm6ts w@ .pprexi@lely of 1,2 tb dd 1.4 tb in siz., slighdy m.lldthe llc CLCUD DNA hr8ndls. The diflcm@ h the siz€ of DNA A conDondts$88catql th.t rlr* rwo vitu6 wft dillcM| Th@ !tudio cl.dly sugaBt d tbar

Cr-crv in Nodl An6ie &d CLcuDh Asi. .!d A&io w@ d@ ro rwo disdtrrGtuidivitrs. Sie whicniB tr&qil ln. dise, d<l rhc ,rnprom w@ indiqliwofgditrivitu bfaliolr a bcgonoviu ws &ught 6 the ca$.I agdt ofttc dis.&.

In 1993, r oompeniE beguovitu! w$ foud in Cl,CuD,irf4l.<t @noD pLDr.,dd ws nMar @non lef cul viru (Ma|t&d a zJ., !93). Howaa, hnfhgirilfalio!! do6 of q,ciD w@ u..bt. to irdE rhc .hlsLristic s'urloc ofCrcuD in conon; th@forc, 0l!o fetoE irvolved in bducing CLCuD w@ q.luai€tt.M,@or .r .l (199) foud r sDIq $DNA noleul., rm.d DNA I rh.r E@iaIod with CL!D. DNA I @uld *f-nplicalc ir pt ,t cdk bul Equi&d Crc[D &!

*tad in plsls &d fo. iD!&r r?rmisi@ How6, ir B fouid rhar DNA t pLys nopart in inducing disc E nploc. Refrrly m addiliond nall ssDNAc, l35O

rucl6td.! mloule rmcd DNA I ws ilold€d nom Clculi!&.ted coton pl&1s(Bndo! ,, dr, 200t), DNA I EqunB CtauD for r@licllion ald 4oluLria .rdwh6 @i@d4.d Mr[ CIAD b c<,trotr ptsG, ilrdecd ,y!!4,totu qpic.l of CrcuD,ftu!, rh. b.gonoviru! (crCuD) DNA p @bprd rcptldk thc infedous uit

2!

.,q!rp!9!.'-......-. ...........-.!9.'l!{sr..!it9$sF,

Gposible for crauD (B.iddon d al, 2001). Fivc viru€s lhl(ed with lhe ctauD@rnpld on rhe hdia s$olrinh! h.vc ba id@tifi€d b be lMoDsti&Bcg(@vitrB belonging 1o aau B.g@iru, .rd aoily Ctuidvidd.c, Tbe sat lic(DNA p) ed sdenitclilc @NA t) @DF!mt5 h^r,€r ro b. clasifi.4,ulough r[.DNA I @rnpon* @ clo*ly Elncd lo,.!d rbouSh ro bave digilat d tDDcoopotr@t' of a s@Dd greup of silgtcsl'.dd.d DNA vi.!s.s, rhe tuvitls tbn r!.

Ih€ begomoviMd sssi.r.d wirh CtIuD, likc all g@iniviru6, have 1wid.d

8minated p.nicl€s, appmxioar.ty 18-20 m in die.rer dd 30 m lMg @strirS oftwo in@rpl€L T = I ic@al|ds joincd toScihd in . shlctuE wilh 22 pobmeric

crpmd8 aad 110 idolical prereir lrbMitr. It is pbbabte, atlhough not ercluivclyplov4, rlal th€ DNA I ed DN,{ t @mporeds, b.ing half rhe size of rhe lnd.onpoM! e 4cap6idar.d i. Ddbhdic, 6tha rhs cdidsle particl6. M&Mr dt

dr. (1993) effm.d th. .lsidion of . cminivitu wirh ih. laf ut di!.e bypolyn@ ctai! re.tior eCR) Mptifqlion ed 3.q!6@ erltsis This E rtlc f61Eponplblis[cd mlt .bhctdiz.ridof tt viro

Wlilc atudyilg th. nolwul[ .sIeG of @non laf dmpL viru (ongiD.rilg

nln Non[ Anqica) ud @tron lef .ut viM (oi8ilrlinc aon parisr,n). Nad€.i!. ,raa, (194 foud rhar tne vituld w@ onty di.r..rly rctat€d Miu 0992) ahdpr.dsftlogical iddtif@tior of th. viru by Bing bd@lor.l etibodiB prcp{.rt 6ohIndiD c6sva nosic vic llreu8h T S ELISA. Th. Esults of l[€ @loeical t6t!coffoed th€ pBw. of a Chilivi@ in infccr.d coiion, Khdid 0994) rerbned tncr

CLaUD panicb weE ob@ed by imuosrb&l €l6hon nim&opy &d alsd.t*t€d d. viG by ICR ir ptdt ti$uc ed indiyiduat virulifftc whiletv (rezi,rl

n&ilon e, "r

(1994 d*ib.d mcthods for d.tedlg rhe riru by ELISA with

bdelonal dtibodid (M.Ab) &d by rcR ft.y de u!.d the rds to ditr.MrisreCLCUDPK tDo 6ve oLtd whir.flrrrllmncd Cdiri vi|re GEitrs ir vMoucrcps .nd w!€d sp6i6 i! ?.tidd .d fosld U.t n hrd d@ .trdriB wirh C@i!iviru i&lal6 ton ! &w oihd stcci6, mtalty otsE patisrrni ftp6 @ inf@l€d sirb .

u

-c.F..p!g.'....- .._- Rdi*ofutmt!&

mtc ofdistingujllte! blt Etalivety ctoscty rcl.tcd ehjt fly-treMined C€dili viru$!.Som. ofwhich 6dhl. rh6e found i. tndia. zhou .r .1. 0 98) nad€ @mDleE DNA_A&quac6 orninc ?a}jrtani cmini viru! hobr., &on r€af Nl affeted coiton (CLCuD-P& or Aoo oka, ed the prltid 3.qu.nccs of weral attdirional isotai6 w@tttldmined. S.quocs of isollI6 noh conor w@ of fow t ?6; i$lar6 ton taf culrlT@r.d oln hd virtudy ln. @e !.quft.. & rho* Aob @rion, I$lara tDh FIoeE n n@ic.fier.d ola! w@ of rso lyF (OYVMV rype 2Ot &d 3OD, bortr dirrirctfibn @i orhr bu1 clo*ly elared ro rhe viru i$lst6 foh @.u.

Thc conon led curl @i.tadc in p.Iirls is €us.d by sddEl dietinci v&ists,wuh @onbinalion €venb involving OYVMV sd oth* unsDsified cdini viro6h.vi.8 !rcbably bd hvolv.<t in their dolutior.

2.6. hl€rltre of rgmm! c .!d tb.r qu.lty rnitl, .!d O.tr Doslbt.

Ahhough the wo* otr tnc inp@@dt of 6n@ Dtdt rs staned b.IoE thc

@rd€d hisrory of culrivatcd crepr lh. |!3.eh bem. largd{i@r.d in rh. bcgimingof th. l9s @tury. -the efdr! wft @lcnrcd duing ([c 5G, and bre.dirg srrarcgi*wol on chegine keeinS in vi* rh. n..ds of inc!6ing popul.rior Th. siolifc rcrtnadc oG thc y6 hs 6ult€d in rh. dd.topndr of a luge bo.ty of g.rElicinfomarion or djfi€mt planr lrlits of @mhi. inr@st utut€r v.rying ovimMdr.l@ndidom, IE nl. of vdiou oihd pldi .h@t .is1i6 Im€ly aMbd of bolk Erpl4i, linl p€rcolage, boll weight, nbq rir.n8i4 bqkine togr!, eglc of ftuiiingbr.Dch, pldt h.idt, nuirirS b.ech trmbd ed pGiiion of 6nt tuiting b.ech ddili.ir .5siatiou wiil @n olha phltd d inpofi&r rele in l[€ inprcvdot of ihc@notr pl&l (Che od Ae,t99l). Th. grcatBt dnEl.f..rs of vEiou r6iis on ,..d@noi i.ld ptuvided euiddiG ro trEdGE for nrlira &t-ri@ of l[. d6ir.d Dtal!Gond a al, 1992). The eFotu &e by U. brc.d6 duira 0E pasr rhE d6.dc hrdbFught . si8Di6@t bPtrhrclgh i! .ouon Foducrion i! !fida.. How€vq, rh.chdSir8 ciEuDstm.6 in lh. wort.t neasirlr. rh. bree<td tur mrinlcd ed.ticimPbvanot in pltut by uiilizing rh€ pot iat 8.n tic tue6 availabte.

.qhP!9|,-.... ...... ....... .......!pti!!rer,.l:lt:.!_r.F|r-

Nwtua sludi.! hd q€ted rh. hith6r ed sigljficer positiv. .ssociarion oftmbd of bols Mlh orha @emic trair. ir inDDving ptdt icld i! @non (Digj,1987; TrrAi, 1994 ., b; Ultoa, 2UJ6 ad An d at.,2@7J ws idarifcd in dliq6!din8r. Th6 in@.d pmd61iotr of corion pl&l t ihc eull of mey i .@lioe ..g.rc6ponss of pl&i ro vdioB Linds of bioijc dd a-biotic .nvituMatd slr*s. Thcpo3iiivc €trdl of lef e! ilder on ptst pbdlctiviry ot @non plad tbrcug[ bold.vclopmot n ee !!d iB n g.liE switrion wi[ hry6i ind.x lrd s&.t orioD yicldwa d?ortd by DbopG .nd .talrade (t989), It. oL of aldc yith p6itiE .f6r! &dofrhoe showing ncg.tiv..fIels wcded th. trge potdtiat for ld.prinS taf si* edshcp. !o ft€ reed! of paniculs pnducrion e8im6 (Jieg a at. 20OO), Studio rcg.rdiqi!. ule of gae &rioo coftroUina taf @ { djt!.mt gb*ih ,rag6, 65, 95, t25, l j5ard lE5 daF sna &wing @AS) tu d@ gmq/p€ of ,rddn @lloo, dd dl_pGibrc@way Fr hrtri& *r5 d.rct@ by paril ., at (199). cd€ric an lFis folowins bolhln. Halhd'..ppMh (19s4) dd Giiffr8! .ppI@h (1956) rry€at.d the inDon4ccof g.nd &tng additivcly dd en-additiv.ty on cortrolirg rhc apEsion oftegf @ ai.ll lh. Aoqt} s1!g6. Hop4q, O€ p&lbnid of.oHddirilc vsidid M hida th6additiv. v-irion ar 65 dsF lnq ewjr8 @As).

Six co$6 w€re idotif€d o the bsk of high pe, !e pdfodd.c for l€af es,Ofih6c c@*s CPD-103.2 r MS-53 €xhibiicd hgh lddilivc gd.lic €ffeq suggcriingth. doptiotr of.ioplc sclcriotr pe.d@ for hiSh yielding ptdts. In @drey, orhd nv.cl!6G *nibit d tLc p|lacc of !ieri6@r ro!-.ddirive g@dic .lI.cis, sigifyjng 0,cioponeE. ofintd,nrilg ed bipdrDbt nting st!re8i6 to b..rtopr.<t for inprevilgth. iisib litc leaf'g.ad se.d @fton yi.td, Batoch (2004) ddin d tnc €fl€cr of t.dnodificdion o! dlitr€$, yietd dd fibe. f.jrs in C$rypjld liBrrd L. studlng BCr

ard F, populati@. The !.gFgating populdios w@ ddivcd tle na ibti,6 byinvolving 1a cN @biratic dieilg for only laf .t4. i.e nomat, ot@!. sub.h!drd rupd oha ThG cull3 of rh* !tudiB luga6r€d for rh. r.pted€d ot sb.kr.l€a{ with th. @mal le.vca in @nd io inpov. *€<! cotton i.ld,.dli6s, tongd edoifh fibd slrFdoriry of sub{h! lef @non ova td€ e@al 1..v6. Th* 6trdjr8,s8g6td fi. potential of mlr!trt lav6 for ti.ir Gxploft.rio. i! tuhr otion bGcdin8

26

.-c.!tP!q ?

Nmbd of bolh pd pl&i ir d imponei @Dponor of pldr i.l4 dd@m@ial vei€tiB diffd widcly tsn @h orha for rhi. ch.rcta. Th. ch.furd hld

bd reponcd b h€ conirollcd by $. gcn6 &li!g lddilivcly dd non-additivcly,

iNolvina e.?i.tatic @hpoal. Sn|di6 of Weg.rd P& (1991)aNdM.i-Z ea..t at.

(2005) H€dcd th. pGtu€ of rcn-addiiiv. gd6 .fects, dd in.ontrsr Kdsy ddGlrs (1988); dd D6hp&d€ ard Bais (2003) r[ow.d ihal nmber of bolts vs@nuoled by tte sa6 &tiie adrliiiv.ly. Howwf R.hm& a al (1988); TFsi 0988)ed Men d !l 12003a) reDoncd ttc involvmot of both additive ed non addiriv.

8mai. .fters ir 6nt olling nhba of bolls. tr! &nc pt{l. h.raial hhaik@ ofnhbd of bolls w6 @dition€d by lhe sas showiry overdomi@€ (Sdim er dl,1986). stdi.! ofK.l5y ed cd8 0988), Mdt er at (2003 a), od M.Ctty.t ot. l2Wa b) Evel.d th. involvdat of signficdr .ftets of.dditiw r additirc (AA) e?istaric

cfiets- h addilion, iorddlion due 10 addirive x domindce eoistesis had at$ b6.@ported in ln. Ir.r:re (G€d .r al, 1974). Tn€ iwolvoar ofrhc e*t.si5 nEt6 rhc

!.l6dd ofd6inlL rl&ls ditrcult,

Ih. digni66l cne.1s of soral dd sleifc cmbidry lbiliry (Mqr e, ol,2001a), rcI. of boll nMbn s m6u@mr ro set@t indirEtty rh. m$s having hi8h

donimce .fi6ts of lint yi.ld in conon (Ju , ar, 2OO7), ed o@mce ot h.1@tic.fiftts for nMbr of bolls we n?ot€d by wo.k6 (ArMi er,l , ryr; Ktttsai a at.,

1992i Sohu er.l, 1993), Wh6 lignificdt dout of vdialion is premt in ile ptethdai.L n b@m* h@ wtul io lh. @ton brccdcE to .vab.te thc qrai ofhditability. Role of hi8h hdia.bilily for fi. sletio of rh€ dcaiall€ gftog?e in edlyg.nsatiou (F,. F (B$bag dd Ga.d, 2004) re ale reDorr.d c&lis. tr w$ @aledtar nmbd of bols F. plat had nDderal. ro nodalrcty hjgh hdnalitity (Singh d

"r,l97l). h c6e ollow hditabiliiy, dela'€d scl€crior i! t@mdded, wnillt in cse ofhid hditability &lelion of tbe dsinuc gdoryF gturyp.s io @ly gocrrioG (F,,

F!) wa su8r6r€d (Babq and ccn6, 2004).

Li*c nMbq ofholk, boll w.i8ni b .t$ a najor cmpondt of *€d conon i.td,ard had stiong F irivc Miarior Mth i.td ot...d 6notr Gy.gi, l98Z dd l99a!). tordd to invBligale liie gm€tic n.choism! conhoUi.S bolt w.ighl, nueous sludica

lnd b@ m.dc whict E eld diflqins infomatid. Th. r@rls ofyadav ed y.d.vt

27

0 96?), Xdsy ,nd Gdc 0 988), dd w@g rnd Pd ( 1991) hld intli..r€d lh. p|lsfu. ofaddilive sde elldts for 0E coltrol ofbou weighi, dd i..onha!y, rcle ofdonindcepFparie of g@6 l4 dplaiz.d by R.hnd ald l(|le (1993), Mat 4 dl. (2003r)

ad Mei-Zh.n er al. {2005), In son. @pon! th. ol. of borl addirive &d non-addiriv.

a6.tic .re|s h.d ba adv6.1.d for lh. ioLdrar@ of (his chdctcr (Sind .r ar,l9?D. Itc qork of sdim .r al (1986) lhow.d rh. F€ence of ovd domi@. ettapF .d io be @nplical.d eiih th. involvdai of ePisrlris (Sitva &d Alv6, t9E3;

K6em er dl, 1984; K.hy dd cs& 1988, Bdbag &d cac.r (2004) fobd ihat

6tinal6 of hdiirlility wc oodcn& ro higb MfmiDg rhe Fsibiliti6 of *l6tinsthc d€snlbb gdot}?q frcm rhe segrcgatirg g€n@1io!& F? dd F3 s sugg€siett bySingn d ar (l9l). Howed,ln. plEd. of .pisr.sis 3ug86ls th.l *tdtion fo. tnepluts wirb d4ioblc hon w.ieht is not srnighl fossd.

LEkcn ( 1989) .!d S.hu €r a/, (193) had Epoded fie @culmce oa ha@si3 irboll w€idt, whilsl i, @nire! Salih e! dl (t 9s6) dd ('lasdi e, !r (192) rcpdrd tl,eecm@. of m etul h.t@sis ir dd! chamrq, ro onld to impnv€ boll qcighr

ihrcugh eldtion ard br€.ditrg itrc rcle of signifimi er*r! of g€dqal ed +@jfic.onbininS ability hld bq €opnsized h wort donc by M.n "l,l. (2001.). h lomercpo.l5 low g€ndic dvec had b.a Eponed (SingL and SineL l98l) whiln in othosrudi6 it ws high tge6 et at., 1984), whiln Kne &d Teiq (19s4) Eponcd4odcrEre 6tinar€s ofg€n ric advd.e for bou weidr

Cill ed Singh (1981) hrd chphaizcd rhe liS,iflcecc of bolt di4 jn in@jn8s..d @tl@ yi.fd. Studi6 ofBb.lldc and BhrL OgS3) ad Rard!.w et dl.,l9i6|n nEveded lhal boll siz ws conlrolled by addirivc gdelic cff€cts, whibr Ma€dith &dBddg. 0972) h.d Epon.d d go6 ering lon-ad{titiv.ty in @nloltirg botl !i&.Chothaft a ar (2003) srudy sho*cd thar gd€s wilh a.tdiiive ed .toninance €flectsw@ inponor ft. pl!fu of.pistdic cllets wa Epon.d in en ptdr mt€ri.l byRddh.M er al (1986), while Sinsh r, al (l9so) roud no episLlic crrerr. rlF61in.16 of ll@w s. haitali|i|y r@ hiSha for boll log$; bott wid6 nriG, rhafor boll width (zneg &d Mci, 2oo4).

2a

.qE !s.P

ror studina s6e &tion contFlling sed conon yi.l4 nmqou Btudi6 by Yadav

.d Y.de 0987), klly &d Grs 0988), !"d Tlr8i 0988) shryd tht s&d @non

ield wG @nirollcd by $. rddirivc gmetic .lIecis, while ir othd studie by Bhar.d€ ddBlrle ( 1983), R.je a dl. (199), &d Ddblde ad Baig (2003) 1bc s.€d allotr yi.ld

trdt we repon€d b b. @nirollcd by rh. gd€s with @n addirivc gd.ri. €F6is,

dow{q, mc mrt6 (Prtaeb, 1982; Yingxin .!d X.rgioin& 198; and Abdllr! rral, 1999) repod.d fi€ F6c@ of bol[ additive ad @{ddilive .omrbnob of gfletic

vd@c fd pldr yi€ld. Mccdry a al_ (2004 .) Elott€d dditiv. x sddinvc.?isdic

cffdt! to be involv.d in th€ inldirree ofecd @non yield.

Nm@u |tsdh wo.t6 Kri.nwui {d cullssL! (t9E ), Silv..r.t( l98s), sd Sohu a aa (193) Eponed occuM.. dd €xploit tion of h.r@ti. .frects in

&.d @nor yield. Soo. cls for rh. @u|rc ofhi8hE Lv.tofilbF.didg d.p6io!in wd @tton ield had bd epon.d by Y.dav md yadav 0981, *\d K*^s6 et at.

092). Modd.& io high tairabitity qriDal4 (Khd dd T&iq, t984), !d bigh

€xper.d eo.tic adv&c. (Ked.r 4r, 1984) p6 Fponcd fo.ll|c rnir ofs4d corlo.

Gimint pdml8sc had posiriv. disi efr€ci on s*d @trd icld (cill md Singh,

l98l). PFioq e*6 hd Epon d 6! FsitiE .ssiatiotr of lilt pffiLAa wiihsd.ml trails of @nmi. imporr.M Gom ,r .t, lg2; ed Trqgi l994a), SinilrlyPosiliv. Mialior ol lirt peffir.ge Mth bot siz &d fbd fndds ws !tudied hyw&8 ( 1988) and Ty.gi ( 1994!} hfomilion on dF iDhqirarcc psdd of lilt pddrag.@ r@@u!, but Ihey lppdEd to diffd a4.ordirg b ln€ phrl mat .irl rs.d. Srudia ofCrathm et at. (2003) &d Mdr ., 'r @003.) had @alcd ddirirc ga.lic .rcrqwhild Mccany ar zl. (2004s) h.d mphaizcd 6e iNolvdot of tdd. paMts wjttrhighd lddiliv. g6e Gff-ts Th. Epoi! of Cng .r al (t987), R.tDd .nd Khrn

0988), Lrckct (1989), Sinsh ,l at (1990), dd Men .t at. (2003!) show.d l!.'bpdi.e of mn-additivc gde cflc.1s in lhe e\pr€sion of tinr rftdtlgq Howdd,sersi er 4l 0982) dd Ddphdd€ ./ dt (1984) h6d advool€d fo! the rclc ofbolh.rlditiv. ed Fn-addirivc g@. .tr..rs fd linl lRan!3e In .dditioa M€.rty a dl(2004 a) i ical.d signilicdl oL of aisrali! du. ro of ad,liliv. ! lddirive cenc &rion itrtbe 6ftll of lilr p@or.g.. Ir! eb. ptdr batdiat gd6 snwi!8 borh donit|'e ald

29

ovd rlohi']|I@ clTets, &ted [o @rrr.l lirt pc.6t!8e (s&]s$i et ol., l9a2: dtl

Hctcrc$ fd linl p.rc6t4. do6 Mu in Fr gmdrion, whi.h.d bc qploit dfor inpoving sd @non yield (Phmds d ar, 2003). lr addirioq Mccdy er al(2004a) Epod€d fiG.inild valB b€twd F, dd Fr popul.ri@ ard !@ hid-pcmtvalB for l!€ Blimatior of hei@iic elf*ts. The p|tgce of noddt€ to high mw!4e hditrbility !u8g6l.d thc po$ibiliry of ibDroving lint p€tgr.9. itmugh s.larion&d bedins (Bhailde dd Bh.l., 1984).

Fiba ldgth is s inpottait etunic rlir in upldd co$on, md dctcmin€s Ur€

qurlity of tle lint pFdued. Fiba ldgth is l6ir!bl. in mtut!, ed hid be€n .pon€d, robc atrer.d by difl@r &16; sry.61ddh6 (un ed Zh@, t988; Nad.rqjo DdRecasy' 1990i Wane &d Pq l99l; dd Md-zha .r al, 2005) had clort€d 0Finponec ofadditiye go.ti..'Idts. Studi6 ofRcnrn,, aa (19E8), Nad{ajd and

R&86dy (192), &d Rlje ,r al. (199) rvc.l€d pEo@ of non adtririv€ sde&ri@ Th. Eports of Tl@ (1982), Desphod€

"r 4l (1984) &d cherhd ., sl (2003)

$ow€d ilr.t ga6 with boIh addiriv€ ed nor addiriv. goctic etrels w@ importot in6d.oni!g rte irhdiranc. of fb.i togrh jn conon. Ond e, ar. (19?8) had Eponed rh.lIibG logih wa c@diiioncd by one g€ic, whibr Shot y ?r ar (1986) r.Doned Oe

'nvolvadr of rirc geG ir thc irhdiraft. of 6b6 lmeth, The wort of Shoky er ar

(1986), Nadlajs sd RaS.luy 0990), $d Babag ald cacs (2004) hld Bal.dhid 61im.t6 of hqitalility, sg86ting lnrt rh. d6ircd gdoq?€ my b. sleld indly sgrtgating Sderalio$ lilc F, ed F3. Itc work of r.scarchG (Kalsy ed Vilhal,

l98q AdMv, 1981; Aknundo\ l9'2i AM et at., l98l; dd phurdm., at 2OO3)

rhowed thll helcr@is did ecu in fibd ldgth, which @uld b€ ccnn nom otutic.l

Dq.lopmcnl of conoa vdi.ti6 with fi!. nbd hs b@omc the nosl desinuerail thw days, Th. potarist to inprevc nbd fnd*, rheu8h scl*tior ed bE dirg,bd b6 crorled by Klorh (1998). fte wolk of rrm e, 'l (193) had shown the

n.8.riv. eiarion b€{wa fihd srFngh &d fn.|]s, wbilsr A2Jl8 a at. (gut ttdKhd er ar (1991) rcportcd negdivc saott?ic o@l.rion of s*d codd yi.td wirh fibrc

lagth ed 8na*, How.vd, TFgi (1994|) had Eported potirive g@ot?ic cortlsdot

30

of nbre 6!@6s with lint Frc@t gp md boll sizc. IlE wo* of Lin and ae (1988),

N!&rrjo dd Rar$my 0990 dd 1992) had @e.l.d additiv. sa. etion in

controlling E fih.r find*s tril. On th. contrary non lddiliv. Sadic effats wa€

ihoM by wsg !d Ps (1991), Naddje rd Rqgasdy (1992), Mly dd Gl@(194), Rlju d zr (199), .d cha$e er .r @003) fd @lttollins ll|€ !!it or

fjl€IB in @notr Thc iMlydat of oi6Ltic cfrds of doDi@c. x domi@e (G.d

sr 41., 194), and ar .rldiiive t lddiriv€ lrtur! r9p@ed ro .omplior. thc i.!dita@ offibd nnd€$ (May dd CFi. 1994; ed l&k ?t al, 2005). ltc wo* of B{r538 ddCaca (2004) d&l.d rh. nish hdihbility .sdnat6 for lindds sree€€tilg inal

*letion of ple13 *i0r low midrlirc vdc 6 be &hid.d in F, ud Fr aa<mtios.nF @u!!@ of egaiv. het@lic.fieB for 6be fi!l!6, (Am.l ul, 1983), ed likc

vdc in F2 dd !r popllarioc dd tr hid-penl rdE hrd b@ r.torrcd by Mccarty

h brc.di.g for inpreving conon pbdu@, the r€lue of fibrc matuity .on€spond!

b hieber fibft 6nd* vdue but tbe brc.d.ar choice is alway! fo. hiSld fibr oaluilyv.l!6 al@g wirh lind 6br, vdios g6.1ic atodie @iiollina 6bE mdunry hd b6ei.d oul by dilf.rcnr tq.Nh wort6!, who bld i?on€d th.r .ddilivc (Nad@je ard

Rdelsdn 1992) &d non addiliv6 s@ .fdtg ryr.hatd.r 4l l9E4; Nld@je udRds6my, 1992i dd Mry dd Ct6, 194) prcdoni@tly @nttoUcd fibrc maturity.

Simildly, ihc digcnic non-all€lic initulion with mn-additivc go. .fl@t predminatcd

old additiv. gac &tion (Nad@jd 6d Rdgrsmy, 192). Th. oq.r@c of rcgalive

hd@is fo. dbd n turity had also b6 r.porrcd (A@ ,r a/., 1983).

Krolh 09E) hrd ruseescd lhal libr qualrty llits nay b. inpbt€d by

inc@be flbG lnqgth (Nueh indirel s.l6tion of @all6 pdih.lq. Prrically, lhit

k no1 po$ibL a mdwing pqirn ld not only involv6 the sdditioml costs bul &.lso

p€iineld's afcci8 both 8br ldgih &d shagth, Fibd st@8lh had b.a rcponed to be

@aitoll.d by 8pD.r &ting addiriv.ly ard rcn-rdditivclt .nd al$ by gd6 shwing non-

dlclic efr*L. Li! ed Zh& (1988) ad Ch.'lhm a ar (2003) hrd E on d tnat g66h.vins artdiliv. .ftelq wbild ltlje a al, (1999) had sielif.d th. ole ofNFadditiv.gde aclion rpvdcd fbrc stneiL Ho*dd $ne wo*s ha<l t!! rt d thar holh

dditiv€ sd dohi@c. .IIers of gm.r wct! GIosibl€ for contioling the fb.rlrqEtl (Muts er al., 2004). Ih. lrudid of Mccdry e! 4t (2004.) &d J@L er ar

ll

.+|J!s.,,

O1|0t) llteld lb. hvoll.|t|.d of ddiri!€ , ddilire 6d do6i!.e : dmi@..Dirrrir $d!d to coEDlirrr. sdcdid. lrdiN wr ty M.r.di6 (2@t .[or.dt!r! hinin.E dnib.r of F6 @rouirt fbr! .t@gtt wr| in th. rog. of I to 2,

Ei8h.r .din ter ol h.rit6iliry, F&!* of hot ft6i!, sq€riority ol pd6ts for bltridood|bi!.ti@ .!d id.o66c.tiN of bvounbl. .[.16 b. t!.ir util,|lion i! hqtridizrioo

Gln\ 1982i @d AtLev, l98l; Mat d 41, 200!b d R.ddy d 4r., 2@t) irdic& 6rrii ir po-"b|€ lo iqorr EbE.ldetr uilizlt ttr gdtc !a||t€.dl$L lo otoo

32

MATERTATS aND MErHoDs

For th. DI6dl invdigatiou otr rh" gdclic Ddhdis @nhollinS lhc

iDlditrn@ of CrcriD, 40 @ttd gEottles diffaing in then e.n ric Dalo! w@

collccr.d &om vdi@ Cott@ Ra@h lrEtitutcrc@t6 in Pa*ist !. ADoog rhe.i0Eemtt?$ oalut d al$ i&ludcd nin @non cxotic emorw6 ia NucotN-35B, R.ba

P-279, R.ba B-50, MK-73, (HAR xAtA)n t86, BJA-592, HAR-444-2, SRI l4-?1, &dRcba P-288 s givd in Table-3. DE pEent invcitiglriom w@ cmied oul udd lleD.panmot of?laat Brc.ding dd Gociior Univ€Fity of Agricullure" Faisat abad.

31. S.E.log of @ttor g.@plDr .8.1r @tlo! h.f cul virur di!@e (CrruD)

Ihc respo|B of,O @non SEbO!6 to CI.CUD w@ errlu!1ed udd ficld a*cU a gbs noN ordilioN. IE d@riptioB of lh. .xFiD.rlal m€rhodolog/ for th.

.ss.san of CLuD in fclC srd EJa hoe e denh.d ir stioa 3.1.1 srd 3.1.2.

Th. po&nrirl of ln. pldr Earaial for CrfuD Birtar@ ws ale sludi.d theu8h

3JJ. Asdrell ofolto! g€rDplsn ln t.ld oldtlto!3In otdd to wrlua& lh. tllpols of coiion Slnotypd to the .ono! lcaf cul viru

dise {Cr-CuD), the s*& of 40 conon lenot}les wft planled in 6eld in th@

r.pliqlio$ following rad@iz.d complct bl@k d$i8rr in 2001. nE 6eld had bieh

mhsl CLuD in@rlM load. ft. Flds of eh @tion smt}?e wft s€.dcd in iwo

w3; eb witb 54 meler long with iltn Dv +&e of 75 o nrirtm !.cds wmpldr.d itr @t Iw wirh & iDFr-ow lpFi.g ol 15 cD- Ite Espons of lhc Dl4ts 10

CrcuD s@ as.d by @ddiDg strnpioE ddclopF@i ad dilee ffiily by

D.ling $v.nl obcdatioB at dittdnt pldn dedopD6Ll st gps ie 35, ?0, 105 edl,r0 di'5 ri6 phnrin& Dala d rh. gturtlic Fipong b d. dis@ re eord.d ar

difldt lss.ftd ddgftq uing th. nodif.d dj5.& $alc 0 ro 6 (IaUc3.D

Plwiouly ued by Athre ., zr (2000). 'Ih. med dil.e ntine valu6 m6u!d for 40

@non wiet'* undd 6eld condit'ons.rc dvd 'n

lable 4.L

l3

MNIIJ29MNIIN4T

slgl

CIlrAD d. Mdtclli4 Fd.CIR-{D & Md*.uia Fffi.CIRAD d. Mo.tt li.r,F'me

clMD d. Mo.4.Uit Fa.

cll^D & Mo4.U*r, F@.

CIMD&Mdts !ia,Fe.

clRAD d. MdlFllir. F@ coMiorlb|Edia

.14

qrp!!t3...........-............. ...........!.d.-.!s1+_!rtq!*!+9*

3J.2. A'gmli of cono! lddph.D u!&r gls houc o!dili'..th€ sd! ofthe 40 con n gdobrld wer. al$ pldt€d in pols (3ix 8@di in eh

poo eith 30 cm di@.&r t tt urdd e itEt-pmf gla ho@ @lditioM. ft* Potr

*@ fll€d eith $il ratd ton lle @non dpaimad n.|d !@ Ar 34 laf srag.,

se.dlings w@ thim€d to iwo e.dling! pcr pol. In tobl, lher€ v@ l0 po$ fd @h

@ft@ rE olyp.. Eeb pot w6 fdtiliz.d wiih nit %ms f.rtilid (wa 9 C&ot) on@ .tund dd.ld np watci wc .pplied 1o tlF grewir8 s..dlings, a r.quind duioa lhc

plet developmcni, Duing thc dpdinotation, prcduct@ of vnal ituculu udei gl6s

houcwaqi .jn.donratu..llyinf6i.d@nonpbdrof S-12, dold vdietyreponed

10 b. bisbly N.ptible to laf cul viro diser. (Ali a zl, 1992). lor Epid tremisiotr

of CrcuD b th. h.allhy phaG, dis!.d plot! ofs-12 BiFd w@ pl&cd hcrwMpl6ts of 40 @tion 8@typer udd ltudy, Itr .ddlrioa mdinur minihm rhp@lu..d h@idrty ldcls requitld .t difre€r plsr gre*rb n 96 &odi4 io field onditioa

w@ maintailcd. G@rypic E3pol@ to rh. di&se w@ t@d.d 3 : ( D dr)5 frk@ fqfisl disede synpbn ro appd on ihe pld6, ed (2) dis.@ @tion by mrking svmlobwati@ .t difr€M1 plet dcv.lopoad nAes; 35, 70, 105 ud 140 dats aftd

$eiry uiry O lo 6 di5.& nlilg *de G.!|. !.1).

3,13, AssDAa of @tton g.rhplso Orcug) el|fdlgInitial *letid of p!gr! b urilja in a rcing programc, inrclEd ffiing

of9lin q NLABnll, Murd!358, Muht-588, FB-900, Fq.901, CIM"443, CIM-482,

6d ClM-4?3 ed CIM-I 100 $.r qFar.d lo b. nighly rcrist ir to CrcuD udd ticld

.d g)N hoe Mditi@. Th@ liE vw ag.i! d.lurt.d uodd ela hoeconditios to @nfm tben cistaee io CLCUD Bing e imprcved bottlc rh@t enninghalod (ALhter ,, zr, 2001). ID addition, two s@@tiblc 8ooiyp6, S- 12 dd NUCOTN

35B '* ds pl&I.d. T@ phtrts nm ch gdotypc aft gdned usil8 lnc "bonl.3hd gEning n.lhod" lo !tudy tleir r.s?d6 to bieh CLCID i@dd lods. CniinS

involv€d naling 3 l-2 .m long x 0.1{.2 cm dc€p cul or thc slm ne$lh. !@nd lsrirr.rD& or6-wFknld pler!. A sinil{ cu1 wa nad. on rh. Cr-CuD inloied bIehwilh . aowirS rip ofrtDur 2G25 m lotrg dd&h.d fon rh. .lisas€d plet of Sl2. Th.

corcsponding cut surfsB wcrc bowd iogaid dld ti.d with pdaflm to avoid drying

l5

rht tLL.r.lradlod!

ard io stop an €ni'y, C@ we t!k6 to brirS lt€ @Eerponding cdbim $f4s into

onr4r. Thn sr€m w$ lla pL..d in | 6 cE lonS 161 tub. wnh 2 cm diddd fllcd {irhdistill.d wara. Diulill.d watcr w$ chde.d cv€ry 12-13 noN for 5 dcys. An . 5 drts,

th* iub.s eerc rcmoved dd pldts w@ obsed dlily to d.tcmine if di$etraJMisid wd &hi4ed.

Br!.d on thc prclinin.ry Nlts for CLCUD dieae nline! in 8la hous, n€E

@nditioB and ihrcu8! gnning (TaU.r 4.1 and 4.3), two v{i.liq l.[AB-lll ald

NUCOTN-35B ca&gdiz€d 6 hiShly Giiler .d highly sw€ptihle D.mts, EQacriv.ly

wdc tunls utilizcd in dw€lopndt ofpl&r m.tdial for innairanc€ study.

1.2. D.v.lopmnl oa pl.!t b.lc.hl for rn. btdit !c. !tudi6 ol CLCiDTo .levelop dpdinflhl matqial io u@ in studying the inhc.iid.e nechdisn of

CrcuD, two vai.ti6 NLA,B.lll ed NuCOIN-35B, ohtsir.d Gom th. @tioa

Smplm (S€tion 3.1.1,3.1.2 and 3.1,3) w@ mssed. Culrivd NIAB-lll @eally develoM by Nud* Islitule for Agricultu€ md Biolo$/, Faisdabad

(hki.t n), dd h4 b6 r€ponqd io be bighly Gisranr b 6rm lerfcwl viB by A}!(arand Khm (2002). In rh€ pt€gt gnfting snr(li.s, NrAB-l I I culrivs wd .!a|!.td by lhe

Imc of M-llt. NlAElll is e dly nslwing vdi.ty, shows boll rctentio! rr v€ry

dly nod.s, high i.ldinS, a|d hs dc.llot fbd qu.lity trair._ The ot[q Ddqi,NUCOIN-35B is d *otic cultivu wilh coe. fibd, n.dim fiba ldgrh bul wirh high

Srding p.!@tagc. ft€ two prMk NLAB,lll dd NUCOTN-358 w@ l]tridiz€dduilg th. 2002 cbp s.afu ll&d @ul.rion of dgirEll€ buds o NuCOTN-35B,

aed s r fdal. pu.ni. ws don in fi. dening prior ro cmsin& Eneulal.d buds

w@ @v.rcd with dmall glGsin b.$ 10 avoid ey polld @dshin.rion. hl!. noningpolla % ol-Ld in a p.tri di.b fio6 NIA& M ed dusred on rhc clteutal.d buds.

ud fi€n bu& w@.ovad wirh bags. M*inm nMb€r of pollinarioN wa nsde io

@llei ruftciot qwtity of hrtrid e.d- Ar Daruiry, cM*s w.F picked individuatly

and gM€d to obLin r..&. SoD. Fosecds ud lh.n pent w@ Irom in Cla houc in

Ds@bd 2002, dd the m.inine se€ds w@ rcsfld for crop se@o 2001.

36

q!rp!"ts ........ ... ........Vr.!!d_"1, !.44 !rJ!l'g-&-

Ite rr plere w@ bekcos!.d lo borh lhe pr@ts to develop lle Fr s€€ds ofh x

NUCOIN-35B (BCM) od Fr i NIAEI l l(Bcrr geFltioB. h additioA fl()l6 w@self-polli@tcd 10 obtain slf€d r, gdention *.d need€d for rbc 2003 cop scensrudi6. ft. r..ds of@h rhc six gdEhrioB i.e. NUCOTN-35B (Pr). NrAB-t I I (P:), Fr,

F,, BCN, dd BCN1 p6 divided ir two equrl part! ed u.d 10 cdy out fi.ld sd Blsshouc CLCUD 6ttan c drl@titu.

3,3. A!!diD.!l of g.trotypl. r.ipo!t6 to CLCuD urd.r tl.ld cotrditiont

Th. 3.€ds oflh. sn &nili6 Pr, p2, fr, F:, BCn" dd BCNr wh pldl.d i! fictd indre. EplicdioE uins red@iz.d conpt.le bl@k d6ign, The seds of e&h fditywft *cded in rews of 5.4 n lolg d|d in!! w s?&in8 of75 m, Tlind !..d! Aeedal 45 cm lpd were ewn within l!. ow. Ihft w4 orc row oflr, P, &d Fr aaeFrio!.10 oes of F: g@.ritidl dd 5 res of @h BCN! dd BCN| gd@rioN.

lxc sil i' whch fiir.xFiD6r ws @i.d ou! E sdyrtay loh (JE: 2j:19 sdd, siL .lay Bpeliv.ly). fta $il pH w6 7.9, EC 1.2, ds/m, sAR 2.88, orgdicolrmls 2.3 7q rot l N 0.06 %, qtd hotdilg c@eity 35.5%, bult d6iry L5 g/d ard

Frcsity oI44.5 %. TIE sladlnt agrenmic p@ti6 follow.d w@ prepeti@ of ldd,thimiry of..€<llings md 2-3 dry hoeitrg!. Firsl irigation w6 .ppli.d 4j days aitrpkrling follow€d by 4 con&quot iri8atiotu pirh & inlcN.l of 15 dalr of cmjddinscop rcquimmc, Fatili4 i.e NPK wa atpf€d @ sledard Momad€d dos!. i.€.

(DAP @ 2.5 baC h@te; U..{j l-6 baC hetasre &d poralh (KfOr) @ 70 Kc/h6ide} Apprepiiat pler prcledon nesue w@ ale ucd wh€n requiEd.

T[. 6!ons6 of plers to CLCUD }@ alsBi.d by Mording syDptoD

d*€lopD@Vs)rplom *vciity. D!r! or the g@or$ic r6po4r to ihc dil.4 w6rdded Bing . 0 lo 6 ntirg sal. CrsbL 3.l) .l difIddl plmt rlage 4, 7, lO, 12 dd 15

wek aft€r ddgtrc, 6 sugg6tcd by A$rd e, d (2m2 a), a ncilDd comonly uedby mdy brcd6 in Palisie in €qtu.ring .orlon ptet naLrid.

37

q!P.9.s M.tcri.lt .td Merhods

Trbl. !.1, Cotton l.f (ort vin! d$e tcrrlD, r.rbg !cx.

Rd!g

0 Complctc abhce of sympion!

Thick ning of fw msll scdreEd veins or onlyPles4. of l€f a{ioB on t@ or ts lhs rc.l.av8 of a pldl .id c&€tul obsatio$

Thickding ofsdl sreup ofveirs

nrictoilg ofdl v6ins bur rc culiry of ldv€g

Sa@ v€h thi.kain8 tud ldf.utirg ar dE rop of

S.vcrc v€b ihickdinS ed lar culjrs on hrlf of

S.vfr rei, rbcltug taf crlilg ad ,tultirg oalne Dr4r w'1h @ or les &ul b@i.g

2

J

5

3.4. Alg@t of g.lotyptc Bpols ro CLCID ir ats, bolr oidfttor!Itc s; familid Pr, !r, Fr, Fr, BCx, &d BC& w@ at$ pler.d in t2_irch

dr@eler pok uds rhe ilMlp@f 8lrs hoe duing rhe 2ool_crop ytu The studyee ffii.d out 6ing a !@edw dq.top.d by Triparhi dd Vllfu (2003). At 3-.t lclt$ag. sc.dlinF wft ditut d io fire pbds to. *h pr, p, ant Fr, 3ix ptDr, for Fl sd6vcplaDbforc!.hBCN,&dBCNr in@hporInbtat tbmw@ |5pt !|5of@hnm_s8t 8.tir€ population i.c, Pr, tr, Fr,2?O pl&tl ofF,, dd l5O plrlrs of @n BCN! sdBCN ef,ddion' Tb. €xpdit@r w$ csiq{t out using a complac mdonizcd d6ignwih ild cpliclrions. For ihis studt pr.durion of viral in@ule w6 mj,r.inedlalwlly on irf@r.d @t!on ptdls of z hjghly susgtible vei.ty St2. To pDmoteFslni$ion of CrcuD, di€a*d pl&b of Sn 2 nisd in cag6 witii hu8c poput.tion ofwhii.fly wft kcti b.tw€q test plmis. h addinon, mdinurv midmm td!.6t@ hdhmidity lcvels

'!quir.d rt dir€mt ptsr BbElh saag6 (&coding io Rctd drr.) w@

Iqded al Eiou btflatr (Appadix-rD. Drta or dE Blot$ of .ll 0E gmrp6

l8

1.5. M.$!nNrtol.grclodic.nd 0brc 6dt .a Dt!. y

Data w6 r@ded on ioportant mionic lail! on 10 gaurd.d !Lnr! iDm Pr. P,

ed Fr, 90 pbnr6 ir F:, Md 50 plmE in dch of BCNu ed BCM poputatiom liom eehtplicalioo 4 d€eribed b.low.

Fuly.xpanded leve at los, Its ed 2Os stmrodiat md6 of uin sr.D fo. aIlh. prdrs w@ m*rcd wilh a pon.ble l.if M deid, dd md l€af !M catcul.ted

1.s.2. P.tioL lddn (.n)P.tiolc lmglh olthc sc tuty qped€d tav6, u$d ro 06@ 0E lqf @ in

@tlon 3.5.1, w calcllat d. M6 p€tiolc ldgd i! .&b faimly wd slculared by4aking Fv@l n€4umdls pa ptdr,

3.5.3. SFpodi.l Logrt (cd)

ragh of syDFdial bE&hcs al lod, 15$ ald 2Od rod6 on @h ptdr wsn6u.d i! cm, {d avehg€ stmpodid ldgth v$ catcular€d for uc jn fu.rhd ealrsis.

3,5.4 Nro!.. ol bo[r per ptetIn rolal, rLcE w@ rb@ pick. Nmbd of botk ws cowr.d !r @h of de ttG

pickings, od the loial lmbd of bolk B *pdt.ty Eaintain€d for @h pldr Avd8.lmbd of tElls for .eh fdily in e&h gddrtio! w6 compur..t for galtic @ty!is.

!.5J. S..d coho! yietd p.r pt.ra (8)

S.€d conor @ttdr.d .t pick- 1, pict-2 dd pick-3 vE butk d, ed roral wcight ofs.ql @non of sh pt&r to. @h f.nily w$ w.igh.d uing ih€ ctairi. b.l,rcc.Avmg. pldt i€ld pa .alculat d tor eh g@ddon.

@pto{ .. .. ....................!-4!ed4'-rrl4}.rgh-.9{'

inchd.d dis.e iftiddc., which is rhe nubq of plels witL sylnploN and disws.vdily sing 0 io 6 scal. d€sib.d in Tablc 3. L

l9

Mrt n&udM.tbods

3J.6 Eo[ r.lglt G)Av@g€ boll w€ight of c&h plhl in dE sii fmiliG wa obr.io.d by dividing th€

tolrl e.d @notr i.ld of. pldl by rh. loral lmbq of bolls picked frod U. plrm.

Av@gc bol w.idt in edh Egucarion @ E€d fd gddic drltsi&

Sizc of a bon o€nBn x bMdth) on @h plet in €&h rmily ws m4ucd withrh. hclp ofvemier Calipd, taLirg on tully @tuld botls at @h stDpodial bl@h !ll0$, l5o ed 2Od &de TtE bous w@ D6u.d on.cn plei ed ne boll sia of€&h plel within @h fdily ws olcularcd for gdelic ealysi&

Dry a do @pL of sd @r!on w€igning 50 9@ non @h pler wsgd€d &pant ly wiI[ r sinelc renq ebric giding D&ti&. 1l!. lirr obLb.d ton@h smple w$ w.iS!€d sd lint pwdLg. w6 catcul.led uing ihe louowinS f@utaad thc !v@gc linr p€Eorag€ of 4h f@ily w6 compuled for d.lrris.

x 100w.idr of se.d @don b r tuplc

1.5.9. Flbcr lrngo, Fin.!e!, Mrtuhly url Str.4ttThe four fib.r ch@tdistics w@ m€6u€d uing lh€ nbrc grapn HVI_900. HVI-

900 fibD gnpn ir r @nput.iz.d niSh volhe iErrMo! wfiich Drevjd6 th€@rbpFhosiv. pDlil. of nw 6b... 11 Ee.sic thc dosr iDportarr fi6d c['@lerbii*srch 6 lastl (tu), r1rd8lh (tho@d pouds pd s!q@ inch), 6Dco6s (rglinch),natuity (p@@tagc) ecording r,o ih. Inrmatioet TFding St.ndards. M€d vdu6 ofrhe 6bq cham.rqistics wn cd.ulccd fd gdctic e.tFs,

3.5. St tidic.l tu.lrlhDala on all th. pldl rriits ncM€d fo. .eh fmity ww oalf2€d folloping

d.lFs of wiec r.hdquc (Srel {d Torie, 19197). Sislifi.dc iliff.m@3 dons$c sn godalioa mc@ v6 r6r.d Bing *tEdats by paniti@irg tt 5m of5qrc(Table 3,2), e@ding to th. ocihod givfl by Li0e qd Eils (1978), BinB lh. fomuta

40

Ss=CqYi)'zt:ci'zSS 4u ofsq@ of@mpdiea.r= CohpaiM @flici4!

I.Dl. 1.2. Ccndo$ for prrdddhg ol.uo of4!.u.bodg rir g.ls.dotrr trrooniogo..t @nFrb.!r (Lta& ud B[b. 1978)

CooD.r&d Pr Pr F, FT BCI BCM

lr !sP, I -l 0 O O 0

P'.vrFrtt-2000BCh$BCNT0OOOI_lF?vlBCt0oO2_t-lP'!,FruBC'qElll_l_t_l

3.7. G.tr.r.dor m.rnr Dd v.riuce d.ly!t,Th. pDgtzMe of ga.nrio! n'@ satysis compuiat.d by Dr. H. S. pooni,

scbool of Biological Sci@es, ald Univ.Eiry of BiminSlm (U.K) ws us.4 Mcssd veidc6 of tne iwo paMts, Fr, F:, BCN" Odk co$ of Fr ,o paMt one) dd BCNr

O&k cds of Fr to p@nt two) SdcnlioG u&d i. tiic biomcrric.t @.trris vft€lcular.d d individo.l ptdr bsi! .d p@tcd ovtr r@tiorioE, Th. @ftcio!, ofgoclic compoDot! of go@rion Dcros e p6arld b T.U€ 3.1. A w.igbt€d tstsqle dalysi. rlge.rt d by Malhd md Jirls (1982), ws plrfomed on the ladtionncs @m@c!rg wird rhc sidplGl nod.t using pamcr€r ,m, onty. Fu.rba no<Hs ofDc'!e18 onplei'ty (n4 Ddh, .rc.) wft fincq if te chi_sqused vcluc w6lignific. , 1l!. bd 6ned modcl ws the one whi.h ha<t signific&r €sftEr€s of allptu t a alorg eith m.*ieniticdr chi lquacd vdu., For @h rlii th. highd n@lElu. rEhI ws alMys t ko s pr in rh. nod.t dfting for iEriee NtAB_ l n4gqler valus for l€ef ar.a, periole length, hon !ize, lunbs ofbolls, bott w.ishr, yieldpd pLnt, fibd logtb ad NuCOIN-35B be6 hiSho v.l6 for CLCuV sdqiry@t'oA slDpodiat ldglr Ciinirg od r!D, tiba frq6! fbcr blruity and fibq

.qrPtsr.9 M.tdid. md Merhorb

T.bk 3.1. Co.md.br. of Sdrdc .tran tor wcidred le.rr rqurc r!.t}!i, ofg.o.ndd |tro. (M.r!d ud Jbtr | 981.

ConFrdl! ot g.!dk .r.6Itl UI 0lIdt lhl

BCo -0.5

.1.0

0.5

1.0

0.0

0.0

l_0

1.00

1.00

0.00

0_00

0,25

o.25

0,00

0,00

0,00

0,m

0,25

0.00

0.00

1.00

0.25

0.25

0.25

0,0

o0 0.5

0.5

0.5

A w€ightcd l..st sq!& s[t]sis of wjecq deib.<t by M.rhd &d Ji.t(1982), wa pdfoded on d.ra fiDn ih. ,ix gcnmtiond (Dd@c, Fr, Fr, BCu .nd BCN|)Th. @lfici@t! of sactic @nF!ars, addiriE (D), dord@ (H), ci(s producr ofaddidle ald dolni@r cfiels (!) ad ovircnFat t vdillio! (E) e prs.nt€d in Tabtc1,4. V&ioB Dodcts wft E, D drd E, D, H sit q D, F rnd E sd D, H. F ed E wd6ttq! ftG b6t Dodet ws st.cr.d, wha & 6ao.tc of Ch sql@ w6 n@jigrifdrfor ![ signific&r p&dtr!16.

42

qrP!:r,g Mrt d.lt ddl|dhodt

T.bL 3,4. CaltLlot of 8fl.tic vrt !e @npolob tor tn. r.tgtr.d t !t jqlmtunylB or g.n.r.llo! v.rl.!ar {Mrl}.r .rrl JilLn | 982)

Corpo.dr. of g...ak nrlrlqD II F

Bctu

BCm

0.00

0.00

0_00

0.50

0,25

0.25

0,00

0,00

0.00

0,25

0.25

0.25

0,00

0,00

0.00

0,00

-0.5

0,50

3.E, E3tlrb.do! of nddori! rrd trbr..dbg d.pr.3rio!Magniftrd. of[€crcsis i' Fr (HFr) dd inh..dins d€pr6n@ u

s€d @tkh yicld .nd ir5 @npon@rq md fibd rnir. ,@ 6u@giv.tr by M.thn &d .tinks 0 982).

E_-

t=

E-

Her@sis (HF,)= E -p,

fic D@ rrlE ofFr, &d

lh. b€ti.r paqi h@ vahc

F, = DGe valuc of F:

Th. siglilicee of n ttuis .d jnMil8 d@Bid wc r6l.d by c.t@LtiDgthe cntiql diEd.n e (cd) ubs l[€ followioS fonul&

InbGcdins dcpBbr = I- r,

43

Mrr.rLL .!d Mdhod!

cd= S.E. x r

S.E. is sleddil ftr of th. diFrtu. of vei.rd n.G, and i! .qunl lo (2MSdr)'2.

MSe, is & ed n@ squr., &dr, ie ih€ rmba ofr.plicdioN

1.9. E.dulior of doDi!u..Estimtcs ofdomindcc (hp) w@ ohrain.d for c&h traii u6ing potcnce 6tio ncllod

(cri6n& 1950).

the tlodimc. calinaic which h6G th. degr€. of dominece of @h ttlit

Thc nid-pd6r v.lu., &d

the m.e of thc ereater pamr

3.10. E*iDdor olrrov 6.!* h..ttiDlllty ud 8.!edc.dvd..Esrihat6 of n@w-tu hrirabitity Olm) wft @hpur.d urilizi's rh. rdidce

or F2 populadon &d sumed vdi@. of rhe b&kcGs populatiotu, fotlMing Wm€i( | 952), fmda sivo bcloe.

Hp

E

l,tr

P

.r - 2o'F, -o'Bc. -d'Bc,

6rF, = vsimcc ofl: gBndafioq

JBC] = v&iae ofBcr popuraloi i.c. (vJhGFj BCr = BCN!) e4orBcf vdim. BC, lopulation i.c, (Whd., BC, - BCN'Sig ficm. ofhdir.biliiy 6inrt6 B !6rcd by h qpreximarcd iF-r6r,on rh.linaeldion dong thc vdiacB oI Fr, BCr, dd BC, (Lieg ar dl, 1968).

Chpre3 Mrtc.trL &d M.thodr

{o'Bc, + o'Bc,,

dfi - df(F:),

The c.b'nat d F v.lu€ ys r6rcd aginst df, &d df:,

Erpet d sfttic .dvecc in rh. tr xl 86q.don w6 conpd.d by ihc folowir€ foduls(Falconq .nd Metay, 1996).

c.A.=K. 6.P. hr

K-slotiondif.r.ntiat, b.ing2.06 ar 5 % hvel ofseteridi @ity, ed6p- rred.d ddinid ofllc ph@qd. vdac ofd. poput.tion 6dd &l6rjonhz - tstimte ol[:m ir 6r4tior of0rc rrail udd cMy.

l.ll. Co[Iordb of y.riD.6

uftrFiclo;,. phdorypi. 16; | tud mvircmold ld.:) ,4iM*" *eEc.lcullr€d iod a..l'sis of vdje6 rrblq

DF MS EMS

MS:

MSr

6: +6;

6:12

r1

DF = dcer..r of Acdoal MS = ma !ql@ Etu.; EMS - .Dr m@ sque dE

45

6i -MS:-MS, h

al =Ms/

Sredrd dM of ilc e@ttpic wi.lg re coryu.d by ttc brdoL 8ivd by

Indllp P, 41 0985).

s3.(6:)-

C =@6cioroflt cdpo@ailltlqFlldE o.qur6,Ir,ffti =n.6.qu.l!forrt i'td!eddf = d.sl!6 oftEldon fo! l[. id F.it

!.12, Coi.hdor ||rFTL cftlalioi! dmg.ll th. pld d drdr€d udd 6cld @ditiotu r@ coqlll.d

rli!8ooquldiz.donsdsl$S&0fdvido*rAdvr4 Shlidlic., 1994.

!d = coYo/ {8" , vrt

= drlldi.d taqa I ed b ttib=@vrtum.t twaEfiltrt'.ib

E edvr. = wiffi of. dd b tri!, E+.div.lt

Msi$,+2

as

6

RESULTS

dl. Vrd.do! ln g.nob,pic rqporstootto! L.fcu vlm3 di5*. (CLCTD)

Dis6c incidqc. of th. 40 .otlon 8.mtyp6 md evqity of dis6. symptong

w@ a4orded foUowire Atltar., dl (2000), Itc D@ g@olypi. Bpots vahs ioCLCUD e Siva in Tablc 4.1. It. rua$ of db.e r@i$ion uoda ficld and Clas

bow sludio w6 100 %. Th€ €slimrtes on dis@ $vdity ud€r fcld ed els hoe@ndilioB wd. vay sihilr, (Tabl€ 4.1), lugeeting sinildily in pr.val@ of virrl

in@ulm undq fi. two diff*nt @dirio@. The ulr3is ol v&ie@ to CrcuD for thc

40 cotlor wi€tid B..lGd hiehly sigDificdr e6oty!i. ditlclw6 (P < 0.01, TlhL 4.2).

Ma vdud p@oted in Tlble 4- I show NIAB- l I I wilh the l.st discs wdiry raring

of0.06 ed O0 uadd glehoM .nd feld @nditiotu, '!+@riv.ly.

NIABI t I cxlibn€d

high shbn€e !o CLCuD, fotlowed by CIMn lOO, NIAB,3sE, FH-900, FH,9O] dd FH-

945, Ii @ftrsl, oths 6 v&i€ri6 S-t 2, NIAB-26, SRI-F4?r, NuCOTN-35B, S- 14, M(-73 ed BJA-592 nad high dis6e ntings tursi4 5.?-6.0- lb.e vdieri€s w€rc nignly

sus&?riblc ro ctauD ud.f boitr gbwing @ditioN,The highcst m€d latdr pdiod g.aicr iln 35 dals ws qhibir.d by NIAB-I I t,

@dd 8bs how ed Mtual f.ld @ditioe Odd onotr lir6, wir! ldgd lat rlpedod ircludcd NIAB-358 wilh 28 ed 29, ClM-l l0O wilh 25 dd 28. Mutanr588 wilh25 dd 27, ud Mutal-388 wid! 24 ard 28 daF in gla how &d t.ld @ndirio6r$pstively. ft€ long{ latml paiod in rh6. vdi.li6 indi.ar.s high 6istrc. lo

CrcuD. C,@O!.s, SRI-F+? I, Sn2, NIAB-84 NUCOTN-35B, S,t 4, MK-?3, $d BrA-

592 hd &e low6t n€e latmt pdiod of 14 ed 15, 15 od 14, 15 ud 14, 14 &d t5, 15

&d 14, 14 dtl 16 &d 14 otl 16 drys 6p6tivcly, $sg6tina w.ptibitity ro @tror

Th. r@6 of ghning wa lm % in al cs. Pleb 6oh ih.2 rw@tiblcvdieti6, of S-12 ed NuCOIN- 358 !4 tlr. frn lo poduc. slrplom 14 d!t6of rfidg6ni.8. Stmplom stut d with mrall vcir rhi€tcning, dd thd prcgru!€d !o s@

!ei! ll|i.kdjrei, upwld aniyo. doMwrd culing of leve *ith v{iou tt!€ d si6of @arioDs, udrinS in plant ltuling 20 days .nd of ineulation, R6*tdt @t1o!

@6ioBr CIM-443, CIM-482, .!d CIM4?3 plots pFduc€d di&$e strptom l?-19

da)s ana gFnirg (T.hb a.3). Di!a5.d pls13 pmduc.d thickdine of th.U so(p or

vcic dd tution 70 d.]s tffd gniiing (DACI Die6e daclopnat or vei.lies

n4.1, FH-900 od FH-901 t@t 24 and 25 d.ys, E petivev Oily two plet! ofFg-

900 dd three pldts of Ftl-901 show€d b.u.i eqnss to ihe diese a tlFy prcduc.d

thickding of fd mall*oneid v.iB, whil€ th€ @iring pl&'8 nom ih6. cultivG

dbw€d Ihicl@ing of sdal greup of wiG. ln @fisl, di!.e s]npio8 on NIAB-I I I,

Mut!*-358, Mut6r-588 dd CIM-I 100 alp.{ftd aft€r 2?-35 drys. With fFt lynpion.

r9pdi!8 35 DAG on NIABI I I, 30 DAG 6 Mubrr 35E dd 29 DAG on Mul!! 588.

Al tne phk nDm $* fow .ottd veidi6nin€s ,ppd 10 h.v. hieh 6tur.nc. to

Ct CUD 3irce 1b.y produc.d fcw snall sstld vci$ ihicLding

Nirc @tton vqi.li.n NIAB.l I I, Muont-358, Mui.d('588, FH-900, FH-901, CIM-

443, CM-482, dd CIM473 dd ClM-1100 turthd ehowed th.ir nighly rcsklancc

Bpote to Cl,CuD uds thc gnfting studi.s. Ulda the sm. eranirg 6tudi6 2

v&idi.s, S-l2 @d NUCOTN- 358 .xhlircd le.rribilily ro CLCuD u<td rwo ditrdar

sowins cordilioN 6 prAdled in (Tabl. 4.4).

Bar.d m lhc prclinin.ry Eults fo. CrcuD disc& ralings in gla house, 6eld

condiliotu dd thrcugh gEnirg (Tabl6 4,1 dd 4,1), two vdicti6 NIAB-lll ddNUCOIN-3sB vft cal.goriz.d a Ligbly Gistani rld higlly sGcepdblc p@1s,

4.2. AsdEdt of llt g@.rrtloB for CtfuD Eirllc ud ot!.r pht tr.il!Ih. ntu gmg?ic Bpons of rhc six g@@tioB ro CLC\D, &d orld

€bnonic dd 6b€r ch.fut rbtica @ tecnred in Figes 4,1 io 4,13 dd T3tle 4.4.1.

An lFis of widc. t@tdqu. w6 dplorld to a!.lrze lhc drr! of lhc chd.t6n6uql in lh. six 8o@1ioro, ard n€fl squc obiaircd e pcated in TabL 4.5.

Sigincdt difrdccc in ncs qua du! io six go@tiou wcr f!flbd pdtitioncd

into vdious intr&lion @DFnors Th. Dd sq!,B of all rh. .he.rd @ aleslltmeiz.d in Tabl. 4.5.

48

M@ diFe *v€r y to cr-cuD for NucoIN-35B (?t &d NL{B{ I I (?' ud€rfelC co itioru w.rc slout 5.51 ard 0,73. 6p6tively thrcuehout thc cDp B.@n at all

Elilgs (ligw 4.1). Si8nificol diEtud (P < 0,05) in m@ glnotypic cpols io

CLUD i! F , Fr, BCN! ed BCNr of0.866, 1.700,2-t80.!d 1.013 E+etivcly (Figre

4.1) e@ oblaind. Sidild m@ dis& wdity tBpotg w@ @r.d uda Cla3rhoe

conditioB (Ttbl. 4,4.1). P&titiodng of Smtypic veim€s al$ cv.al.d tjg ysielifidi dilf.rcrce (P < 0.01) in tlE as.nt!, bekcNs6, dd intcrmiion conpotrqts

du€ ro P'3(Pr x Pz)vrPrrdP',s"rr(Pami3x F)$BC'8x Fr, (T6lc 4.5).

Conpeisn of m@ v.lG fo. lof G prs@&d i! Fige 4,2 d6led lhat

lieb6t ma laf '@

vdue of224.27 @'z w ir Fr aDd low6r lrluc of lOE.5 cdz motffied b NUCOTN-35B. A!aly!i! of wi&e Fv.rkd nidy ri8rufical (P < 0.01)

difiMi oong lh. rix g.t@tiou (T8blc 4,5). Fulha partitionbS of m@ sqlffiofsix ga€r.riore lhowqd lhar i !@tion @nponat!; d@10(Pr x Pr), ed pe@t' (P's

x F,) wft dl$ hiShly sisliicet (P < 0.01) for lcaf ag (Iau€ 4.5).

tualrsb of v{iece for p€liole l6gd' Ev.dcd hichly siifcet difilM6 (P <

0.01) ddg rhc 3ir gddili@ GrU. 4.5). thc inteElid @mporats daltsis

b<iw@ p.Mt! (Pr x Pt, paEls *il[ Ft (Ph x Fr) ale salcd hi8hly sicnif@tly

dif€Mcs (P < 0,01) @ory lld who wd. t6:t€d. Th€ tuniid vaie@ conpondt

dalrsis duc to int r&tion b€rwM two bekcrcsd (BCN! x aCNr) w$ .ducad to

siificet (P < 0.05). Th€ highel Ftiol. lclgrh of 15.3 m w6 obad.d in ihe Ir, ddshcr6-t vcl@ of ?.E4 ch in qotic culivu NUCOT-35B (F,gd 4.3).

IE Do vclE fd ln. sFpodbl ladn in NUCOTN-35B ed NIAB-ll I wft16.08 m &d 10.98 .rr E!€lively. 1l!. Do syDtodial ldgh i! F, ea 16,75 cnr i!BCN| dd BCu vfr 12.02 ad 13.0 c4 rqlcctivcl, Pdtitiding ofmd 3ql!6 (du.

to 8@dtioB), p.lted rhat id.ra.ti@ duc ro Pl: P' ad Bc'! t F: v@ also hiehly

siiner e < 0.01) for BFpodal logh (Ta!b a.t). h.lysis of vei&c. of dE dlla

otr srrpodial Lngtl rbowed illt rh. two plmts, FL Fr, BCM, sd BCNr dilldsisd6@dy for rhi3 ch!rurr (I6le 45), &d rhe diEere|G & cldly d@ict€d i!

DaL o. nMbd of boU5 pc6Ld in Fig,45 cldly slow ! 8rt31@nF8sl in thd

p&d&, Fr, F, sd bekdcss, and th* dilfdtu.3 w@ r4ord.d io h. highly

49

.+|''.!er4......... ,........ B*!c

3isni66r (P < o.ol, Tdblc 4.5). NMbd of bolk also appd.d to diff6 tiSDifican{y C< 0.05) wh6 inrer&tion due 10 Pr x Pr, {td !'s ! Fr, &d BCN" t BCM w6 sMi.d(Irblc 4,5), Th. hieh6t nmbd of73.4 holk es @ulcd in thc fr (Frelrc 4.t.

Th€ nd bol wcidt val@ n@ucd in dx @non gpr@rios @ prestcd in

Fisus 46. Sirnifi@l diffd@6 *@ .,b!a.d uona aa.Erio6 G.bI.4.5). D!l!

tunhd 6bow.d ihat bou weicht w6 hidny lienifi@l (P 3 0.01) wha intcBdion duc !o

rd@G (P, x Pr), P's x F, w@ .tudicd. hta!.tid Edtinc t@ P's, F, x BCt, F, w6al$ obswcd dignifi.et, (P < 0.01).

Ihe n6 valB fo! thc boU size iD NUCOIN-3s8 rrd NIABI l I wft 14.05 dd10.98 mr, r!.pativ€ly. 'Ih. md boll si& in F? ws 12,89 cm': ed in BCNr a t BCN.

wa! 12.46 sd 11.02. cn:, 6!.ctivdy.'ft@ m6u!€d n m valu.! in six cot@

eddtioB e plE@&d in Figw a.?. Sigdi6@1 diftmd w@ obed€d 'mnggcn@tios {Table 4.5). M4 squ€s (Ta!l€ 4.5) show thai boll sift difdrd

ligrif@ily (P< 0.01) h Oe six gdailioa how.q, the irro:.tion *.ooded wirh pr

w Pr, w@ not significei e:0,05). Th. i q!4tiordu.io!'!xFLBCN!xBCN1,BC's

t Fr, od Pt, Frx BC'c Fr, }@ dt big y dgd6@r (ps 0.01),

50

Irtle 4J. Conp.r&o! for tt. rupol'.| of cotoDr 'rrldLdtbB lo ono! l..r.crlvln. dil.& (CLC.D)

Uldc d! LoR oldrdoB U!d6 ubrrr d.ld oldtdod

MNHJ29

23I3

X

t3

t,lt

t.l3

rlTirc t t.n for nd dw $r.sn@

5l

I.tle 4.2- Ar.dydr of v.rb!.. of nth8. of 40 @tro! g.!otl"q for CLC.D !!d.r

So.cofy.rlibr dt Suofrq.G ltr S{..E F-ndo

RAficatio6 2 4n 236 39.39

OcootJ,p6 39 33E.03 Eo67 14129..

Etu 7a a69 0060

Nd-lddilvity I 0.21 0.210

RAidul 77 4.4 0.05E

" - ntnly ristm..i .t I < 1,.01

52

cEP.gr

I.DL 4J: Rrrd!8 ud 8ro!pi!g of @n ! gc!otr"6 o! rt. brh oro.n r6pocto clcrD

Mar vd€nlt tukd or{sr DiIera.dos.l1

sl,gl

MNIiI29

FH6],I

. tSDdp<0.05.bno.Ldrs*idstuitxv.tB&Mrigdf*jd.rp<o.oj

.,c}!r!g-4,........,....- Rdr'

T.bh a.r[ R.I'od!.. of@lor !.rorrF ro Cr,C!D OMlr .rnddd gr.ttlS

av.htcorh cnnht Dra. Jor-"F r,r..r -5;i;. -".;;fr,)'ffi ffi eF. ffH u@E

"d..

'tii'Enit' - dtYt'{

r32t?E

rm J0

]Er

; @ -,,F.'.. " e,ad;;EE-E *ffi ;E;ll' rou otg'oprhr (dim) K drH dn .8" ind. obtd.n

v

I rI| !_&E 8Al.ggrl:t IlE--,1t;! d

t:a E

ts 5.- -Fi5e;.4- z

aFt f; g'-E

!riqE!EFa;t j:Et:^.: a; ,";37r aF"';-Fi-

1:; 6 3 r . g ; I S p S l

=1q:qn$aAla:Fte:-3!:iIR,sa

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!:-i81$!es.is:-i:-':;:+;I.;:t

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rie

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rB@4.1. Md vds orcra'rD diFc *€i9 otrjr dd t6.diNf I

100 r

250

200

t50

t00

FliG 4.2. M.e vdu6 for laf e of in ..tro! t Fdi6l4d: P'= NucoTN-r54 P: -Nr BrrI, F,- f Fili.t c!!dio|l F - s.oold Fittrtr.'dia Bcr=F x NUCOTTN.I,B! ACx'- t' x NIAB -l|| &d rdidrr.t&dld.d m|E'

56

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Filu 4.r. M{ vj6 fd pdiok l6{fi of rir Mo g!ffiB

20

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2 ilPiru r.4. Ma dE fa tnpodid las. of 3ir tu s@dBL.t ldr Pr = NUCOTN-3,B, r: - NI/IB .l r r, fr- rr md eaFdoa F,- sd..d Fili.t a.|ditoBrcr r rr x NucoTN-35B, Bai' - F, x MAB -t | | .ld Edial b|B E.r'd"d.tu m

BCq

51

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Md p€rfomcc of rr, P,, Fr, Fr, BCN, ed BCN| for seed @tlon yi.ld epr.!4tcd h Figw 4.8. The dif€r€@s d. sirikine dde lh. g.nqatio$ for m@ *ed@ttotr icld lhe n6 i.ld of l[. p!mr5 difrald si8nificedy (r< 0.01) yith

NUCOTN-358 havidg . yi.ld of 88.93 I dd NIAEr I r wirh 218,5 s. Sinil& diff.tw6wft obsd€d Moog othd gmtioos, dd rh4. difrdd.d wft highty signiticul(P< 0.01; Table 4,5), M6 squG due ro Scnmliom w@ psnitione<t inio vdious

DterAcrion componmls sd ti@ difr€rd 6ieii6csdy (P < 0.01) for all cmpondt'Md sql!@ Ev..&d hiebly signiic{r (P < 0.01) diff.a!6 eo.8 $. six

gder.rims for gin tm our pd@aage (Tabl. 4.5). Tne inr@rion con4oMrs prvr pr,

Pr v! nr, ud P s! Fr vs BC's, F, wde at$ sisnificet (r<.05). Med conpdison fm sinnn our Frc4laee of thc sn g@.rsriotu 6 pmlEd in FigE 4.9. Ih€ two pMts,NUCOTN-358 ald NIAB- I I I w@ sigtrific&tly difia.Dl (P<0.01) yi0r 41.6 ed 3?.I %

git th ouq BF.tiv.ly. Ih. n@ gjn tm olt p€I€t:g. of Fr, Fr, BCN" wft 3E %

sd 39 % foi BcNl

M@ squ!6 p.rLd Ihat i @tion d@ io P1 vs P: , P's vs !r ed P'3 f1 vg

BC's, F: w@ .bo sigtin@r e < 0.05; Tablc 4.5) for rbe F.it of liba lagth. Dlc b@dba l@grh vdB of six Bo@tios @.hom in Fig@ 4.10, Ite I|r@ di|I.m.6mone ilre gdmlioro w.rc signif@l (P<0.01; Tablc 4,5). Th€ med fibr€ t6grb of Fl

ed NLAB-I I I w@ !imil&, bur thd. difcrcd dicrincdtly (P<0.01) fron thc Dcs ofPr, F,. BCN ed BCN| (Fig 410).

60

2

F{8,1.5. M..! vnE hr Mhs of h.lh oldr dD aerid

c.o..til[t

Ftc a.6 M-! {16 lb. loll wiail ofrir db as|dnL.tDll:P,-N!COTN-354,P:-NIAEntI,F,- l!m adrioqFaSedfilidaFeria.BCh-F x NrcE rN-35B, BCN|. F x NtAa -ll l &d sridl !.rF odni.d|qa

6l

t4

BCNr BCrr

FisGl7 M.s rru6 ror bott riz ot tr m rlldioN

300 .

200

Fum..r Md vrl,6 fd 3..n ffi yi.ld of !r do Sa.Jdrrgan,3 P' . NUCOTN T5B. P: - NL$ .r I l. F,- a Firid i@drd4 Fa S<dd Fili.l s!|drqBCN!= F' i N!corN-r5B, BCN| =F, x Nr^B -rr r ud di.al6s E r

150 l

r00 ]

62

FiCc4.9 r,tdv|lE fesiminsMbof mgtu

E

g

Gne|!tions

risG t.10. Ma vrlG f4 6b.. kng6 .r!n M r.ldi6L.tdrti P,- NuCO rx.35B, P: - NL{ar I I, F,- l. Filin g.ldid! F,= S.od ntilt g.'did, BcN"-F !NUCOIN-35B,BCNr - FL x NrAB-trr Ddv.nic.lbsGMomeltrhm

.q|J!q1

Dab shom in Fiew 4.l r depicl th.r six couon gdmlioro difrerd siificaady(P 3 0.01) fo. nbr. 6]4s. TtG tl@ fibd 6ffi rue€d Aom 1.76 to 5.05. In

addilio4 dl rn€ int reiios sm signG@i (P < 0.05j TaUG 4.5) crcept rh6. dw ro p's

vs f'. Sinil& 6ults wft obrai!.d for fihE m.nufty. FibF nfilitt dlra of th. sixg.oaldoE e shoM in Figw 4.12. n!. m@ fibt mitdry vdB w@ 85.t9 yq

42.46 oA, 84.27 y\ 83.14 7o 80.45 %, for th. Fr, NLAB-ttl, Fr. BCN! ed BCNr.qFcdvclt. HorRd, rh. b@ fibd qrBily of NuCOTN-3'B of 89_60 % Eaf.dcd siSnif@dy frm ,lt th€ tiv. strFrios.

Ard'sis ofv&i&cc offibd 'lj.Dgth

sto$d Itrr 0t ditrG|tt6.noDg lh. six

ecnmtiou w@ highly sisi6@i (p < 0.01) dd inr.Bcrion dE 10 p, vs p2 &d BC,s sFl 9@ al$ sienific&. (P < 0,05; T$le 4,5).

T rE obe€d n|@ wtc ot tibF strEnglb

td ihe sn 8o@tioro rc shoM in Fisft 4. t 3_ Th. md valres of 9j,7? for BCNrdd93.11 FFi fd BCN! (thousnd pouds F rql@ inct) rF siDitd. Ilc tibc srftnsrhof NICOINJsB of 92.09 tppsi ws gFatd ild of 6bq srength NtAB-l l1 by 3,54

IBCr,

IBCu

FisN 4. r r. Mo hlG fft fbrc 6l*m$ of six o!.n mnsL.rb(' ?, . NucorN-lsB! pr - NtA! .|| t. F, . f Fij,,l s*d4 F, $qd Fird sm4bCN! -F' x llrcorrN.JrB, BC\r - rr ' \iAA-l ud Eni6t b6Rssqmmrrl.m

64

.9,t!P9gl

100

90

20

BCM BCNI

FiguF 4. 12. Mro htu.5 rs ftbr rtury ot rr don sdlrrrooe

90

30

20

BCN! BCrr

rig@4lrMuq|u€fqfibm3t4'hk3! L P'-NuCOrN-J5BrPr-N|AB-I j,l,.f f,l,.t s@da F, s.qd rrti.tsdjo!,BcN!- F' x Nu€OrrN.)54 BCN| F, t Nl^B t.iErietborel'lldEdutu

65

.Srrlsl

43. G.n.dio! |tIdr .!d v|tLle .!.trsitTh. knowlcdg€ abour rhc $nft od hagnituda ofg6.1ic cfIat! i! rh. bE dins

mrdi.l sgg6r.d tial elEtion vs ne65&y in odd ro d4clop e agpop.istc

ldano! pbio@l for id.nriftng den€d plsb. I! ord€. io .o[er itrfomrion on ihc

goetic n chais @nhollir8 veiou pler rr.ik of 6dq?iuD ,i6rr!r L., rhe

gdeElion med &alrBis pMeduc outlincd by Mallq &d Jinb (1982) ws u$d. honif !o w rb. bGr,fin d 8@.ric hod.l ro tho dda, de *liDte m.de for rlE sd.ricp.mei6 @npridng difldr cmpotroI! of gm.tai@ m.s @ Siva ia TaU€ 4.6.

llF fou'pgm.ld nodcl d, [d], thl, ud {il |lpqtd ro b. ad€quar. for @non laf cqtviru di!.e. Fivc-pa@ctd nodll n, tdl. til, tjl ard tll dhowed O€ b6l fi1 for lafle It. 6w ptu.rd model @DFisinS orn, tdl, thl, Iil ud [] ws ad.quars &rpltiole lcngth. nMbd ofbotb and s.!d @tr@ lcld data The fvcp&d.t r no&t ofm, tdl, ILl, til.nd tll w4 tlF ber fir for synp.did l6st!, fbd fn@6n ftba dduiry&d fbd !tr@ath, whry eodel coNisling ofD, thl, Iil, Ul, tll arp.a.€d lo b. l[. b.rtlit for bou ste. Th. fou.ptuaa nodel compridng ofn, [d], [i] &d Lil lpp.ed 10 b.adcqu!& for giding pdoragc &d fibq lengrh. It. signific.nr Chi-sqlre (Ir)sugg6l.d tle inadcq@y of th. modct for bolt v.iSh! indicstitrg rhe invotv@ot ofhighs ordd oI g.n. irl@lio! cont&lting thc tdit cxplson.

Th. g€@tion vdiec€s for CLC\D @tion ard orha F.iB e givo in rhe

Appddn L Eslihat s of @npoMts olgadic veies with th.ir 3rldard.hR, sdChi-sqle k1 vatu6 for vsiou corbn pler lair' rrudi€r! e su@ezqt'n Trbt. 4,?.

n. a8l}!is of voiaM for Coetic modets wil[ D ard E, @Dporcnrs gde the b.ri ftfor srmFdial ldgrh, bott ste, !..d corion yict4 giD lu ou1 &d 6bE naiuiryp.Eot g.. fte hod.l D, F 4d E show.d ihe b6t 6r fo. rcetion io ctc\|D, teaf!4r,P.iioL ldgl4 dd fibre lincn6s. Sihitdu for nMbd of bols, bol wcishr, Ubre lhsrh&d trbrc ramglh, the modd wirh D, g a"d E @iDd6rs qhibited rh. tBr tit,

66

{.4. f, imtto! of b.t Mis, hbr..dtn8 d.p€rlotr ud potoc. ndofte 6tinat6 of h.t@is, inb@ding d@@ion dd pot nc. ralio for veios

pldt lr8irs e pr@tcd in TalL 4,8. It. ncgariv. hd@si! ed inbr.€din8 dcp63ionvrl!6 of -4.6? .nd -96-lJ, rcqEtively 6limar.d for CLCuD @tion e lhoh in

lable 4,8. h ddilion, lh. poLnc. 6rio ws .ls !.gative, h.ving a v.tuc of -0.94.

'Ihe bighdt positivc hetercris of 38 Dd 34 ws ob@.d for laf @ and tr@brof bolft. Pottu. r ios for |}* rwo ch@rd w@ 2.1 I ed 3.?2 r+crircty, tr lhe F:

8@eEtioa inbrclditrs dcpE$ioD m l?.E7 ed 28.4 fo. rcd ae ed nmber of boll6.

Glerircly. For plriol. ldgl[ h€t€Nis ws 2,35 in Fr wirh a 20.83 i$r.ldingdepruion obw.d in F goadion,Ite poLnc. !.tio for p.riot. togll wa 1.92.

N.grliv. &d rcn{igni6.e! herturjc .fael of -2,45 w6 ohsded for syrDodi.jl.ngih wifi a signifcddy negativc (P < 0.Ol) b!rc.dine degr.sion 6rimtes of-j.19,The valu. ofpotm. Erio for th€ rlnpodi.t togrt ws 61idcd s 0,04.

A poritive &d @-digtrifcsl h€t @is va.luc ofo.3j and r signjficol inbrc€dilgd@Bion of 0,26 w@ obh€d for hon wighr with pord@ nrio of t.69, Sirdfc&l(l < 0.05) betftsir &d inbrc€diD8 d€pBb! vdler of 0.62 &d L?8, BDetiv.ly w@ob3ded. Vdy high porft. Etio of 266.14 wd 6rimt€d for boll !iz.

A signi6c&1 buI n Sative h.l@sis 6iinde for s€d @$on yi.ld of,l E.47 w6obsdcd, Ch tn. othd hud, inbr..ding d€pl6ion ws sigificmrly !@irirc, ahibitinga value of70.55 wirh a poi.n@ rario ofo.?5, lor gin iw oui ih. lFr.bric efari vaiE of-3.{9 wa signiE.dt eiur posilivc ad signifi@r iDbFeding d.pBior of0.44 ed a

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t

.91'!P94..--.-. RBuu.

Tn n€garive sigrificdr h.{@ric .tl{r of ,0.i3 ed ! riglifcer pGftjvc

inbEding d€lr€$ion of 3,52 wq @oded for nba logiL 1!€ porac. ratio qs 0.64.

A neeadv. liglifkl h.l@is v.lu. of -0,736 w6 obw.d for fbrc fr€l6,ft! ilbFding d@r6don ws 0.094 wirh a lotdcc r.tio of -0.44.

Th. siif.dnt rcg.tiv€ h.rmdc cffet of-4,407 follor€d by lDsirivc aisdfter

'nb@diog depcdion of 0-927 w6 .rijn.tcd for fibE @rurity. A por.n@ 6iio of 4.31

w 6timarcd fo. rhe t ait.

A negal'vc non-siSnincmt h€r.@i6 oI t.58 foltowed by posiiiv. sis ffaiinb@ding d.pcsio! of L63 B d€rl€d for 6bE strogrh. poi.@G niio wa 0,I I.

{.5. CoDpordr' of v.rl.tio!,.!ttD.td ofreor &!e nc.it Ddity {.t 8d.rl

E iiMi6 of saorrpic (6; ), phaorpic G; ) ".d ovieMdd

@ p@r.d in Table 4.9. ra. 6rimt6 for &c csbt pic vdi.M (6: ., for ,tc 13 bairs

tuCefton0.l8-3194.65.

Th. bigh6r valG o13194.65, I t20.03 ,!d | 92.j0 for sed onoa g.td: lefaMdd nubd of bolk pd plmr, Espdtively wcF obsd€d. On lle olhq hdd, 0F lowcsr

vd6 of 0.t8, 0.24 &d l.?E fo l'. geDrypjc v.darc(6;) wft .s:rinarcd fo. dbd

firos, bol w.iaht dd bou ste, cspetiv.ty. For othd t .iis gin lun ou! 6bd toElii,CrcuD, fbd slr.netb, srnpodiat togrh and fibrc matuity, th. gdotric vanec6 (6: )w* 2.42, 2,74, 3.3a, 3.99, 449 &d j.8q Eretiv€ly (Trbl. 4,9). E vircmatrlveimcd (6: ) had n€stisibre inp&r on 1o t6itj i... CrluD t@tion, periol€ logrh,

rynpodiAl lagtll hol sizc, hot_l wcid! gi! in ouf fbrc ldsrt, tibE 6@6i fbEmanuity dd fbd 61r68rh, In @ntisr secd conon i.td, taf g@ &d nmbd ofboll!F pI4r e@ sigd6@dy .frer.d b, avillm@r,

mc 6rimar6 of nmw *ns! h€dhbility of 0.97, 0.s2, 0.82, 0.16,0.75, o.:/j,0.71,.nd 0.62 for gin dh oui, bol lize lef .ra, Ct,cuD, fibrc t 6d5, fibi. m.ruiry,r&d collon yi€ld, md petiole lng0\ rcspsriy€ly w.rc hieh (Tabte 4,9), Nmbd ofbolh

(6:) vdac6

1l

.g'.rae!q,t.-.....-..-.- Ra r.

fE pbat brd I Eodoltc baii.rtilit of o.aa. ft. bdiialilty 6tD.t6 hrcyDpodidLttSlh, boll w.ight, fibe log6, sd f6E srmglh 8r6!rer rhe t,

Thc Gdln.t!! e.d. fo. ttc gaaic dve6 fd 13 rrrits qft in rlE rug. of0,46 82,65 dd e p.@ted h TabL 4.9, Th. hidcn edu6 of ed.tic adve*82.61 58.01, 12.?6 ed ?Jl w@ drn'Gd 6r ..cd @fiod Jiel4 16I .r.., truta ofbollr dd 6ber strclgrb rceeiiv.lt Th. lowcrt vatu., 0.46 ed 0.68 of gendic.trvlB w@ 6rio.rd fo. ftd @rriry sd fba fffi Tbc 6ti-nr Sacricalv@6 for strpodial l6gtb, fibd t@gth, gjn hrm our, p.riotc Lngtt, CtCuD, bolt3i?, ed boll eiSht g@ 2.4a 2?d 3.]& !.99, 4a9 rd 5.E9, G.pcclircty.

72

FF:sElr5:is:x

:33::r353t5Sr

Siss!5s$E!:rR

i!a:X:ri:E:te

l!e:E:r;:r:$i

rgiis*F;3f3F5

e

t

3

lat=

I !-q

IE

.t!

-{95!EJ

zIi

a

E

g

r

I

c

:

e{e9:!!rEiiT}

t

4.6. Comlrdor D.l$L

Th€ ne valu* for el@tiE wi.r6 of six.oton gcn ratioN gFm uder fi.td@nditios I@ u..d ro c.lcuLrc @ctarion @.Ificiots G), ad wft prod.d agaisie&h orhq 10 6how p&dl.ls b€nr€q rl€s *t@tiv€ vriale (Figftr 4_ 14 , 4,25)_

Highly sigtrid{i (P < 0.01) positiv. ..siarion or r = 0.9 w6 obsdcdbetw6 l.af@ dd periole logh (Fige 4.14), i = 0.90 for l€af @ 6d roLl nmbaofbolk (Figw 4.I5.), r - 0.93 for p.tiot. lagrb ed nmbq of bolh (Fig@ 4,l8), ed r= 0.99 b.rwen librc 6n n6s dd fbF natujry(Fise 4.21).

taf @ ed bolt wcishr Fis@ (4.16), rrd lH dd *€d cond lcld (Fis@4.1r, p.tiole ldSth dd boll wlighr (Fi8rc 4,19), p.rioL t.nert dd !Gr<t @r0on yi.td (Figft 4.20), nMhd of bolb {d bot wcidt (Fige a.2l), cin lM out dd crcuD(Fige a.22) ed fbE tind$ .!d Cr-cuD (Figw 4.24) qhibned ,i8'ificdr (p < O,O5)

posilivc alsiaiios of r = 0,87, r = 0.81, r = 0.90, r = 0.84 I = 0.8t, r - O,8t,r!d r =

The 6ulB of @@latior .r.l)Ej! rhow.d rhat nubd of botk had r Dagdnega$v. eiadon of r - -0. r I wnn cta\D e.rion (Figm a.25),

!s|!!!!4 Rdnlrr

Ficn4.la. scerdi.stu forbE tdfe(u1 en Fn b hgrh {q) of'n.fu!ffidB

I@i't.

Fite 4.15. S.e dir3M fo. m6 h.fH(m') ard bo[e rmba of slrfug.dib

IIE.

Fge4.r6 S@dn!@ ft.@ t .Jc(.dt nd b.[ sishr (!) of irdbg.'@am

Firc..r7. s@d&turorM tdf s(.!4 bd ...d .le rLtd (d of

'x.fu3.ldi@

iI

Figw 4 i3. S€@ dngM for pdioh hdSnro, ud tmbe of bo|l of jrijle3dtd

Fhu a.20. scd.r dnsm fdl|la Fdot.|aJn (o) -d ...d d.. ,idd@ orsi: .fu .tuiodr

Fturc 4.19. S.@r di.8m for n!- p.dot.ladr (m) od bou *ti| c) of,t

FigE 4.21- s@ d&n tu 6a bduh€of bol&.d bon Ei.hc)of 3ix tu sdic -

76

Ch.Daaa l-dr.

gi

GO.T

d

Fige 4,24. S.rrr di.gtu 6i tlH fi6c6!.n* !di!d) ud clADofrhtuS.ldiu

FtN 4.23, Sstu diagm for nm 6bEfffi t{ft.h) .!d mtiy(r9) of .le.l|dic

Fisw 1r5. Sd!*d.gm fdllh nmbaof b.Ur .d Cl,cuD of 3n.dh8.diB

n

DIscUssloN

To b.irgine dy aolulion .y chdS? i! a pbr1 cbr@r6, .irhd dreugh nfinztFlation or dirccled brc.dine appD&h6, th. .vailsbitity of lwo cohpon@r& ti61ly,ldation in ihe ch'@id, ud sondly vdialoD mEi be all*Ld by a dg,i66r gd€licconFnot. Epiddic of @rion tef rut viru disc (CLCuD) wd rhc @np.Ii4fa.lor in ile d€cisid ro d.vie ncw 6r6tegid for coirotr bre€dine pegllme in psli.tan

(R€hn{ a,r, 2002), DE CtalD is Miar.d with fou dirtincr b.Somvi@ edtuv€l @mpondsr DNA-I ed DNA.b€t! (Zhou ?, at, l 98; Mlltsr er al., t99a; Sdet al,, 2@0: sd B'id46 .t at,, 2OOl). Mdy m.r[ods cg. ELISA, i@unoldnarelecLon micolcop, ICR ed routh€d-blonins l€.hniqle etc have bc6 ue<t !o d.t ciue prlge or {tMc. of ile vi|rs by mdy r*qrch6 (Mi@., al, t94j x}rli4194; M6lik.r.l, 1995 &d ttusain d a/,, 2OO3), but it b r.poned lh.t the di6se is.or dumtted m*teic.Iy, &d thafoc e i.Ierio6 clon. hs $ fr hetr.@n d(SM er ar, 2000). Thun fcld d.luation is the nGl .fl*riv. add esy me$od !o ue by€otlon brEd6, IE rresdj$ion of CtauD lhrcugh *hileny is mrc &tuat howevd.Dai.bining of a whn ny Fpulatiotr dd t.!9ing dbjol lmpdre hjsn fo. diwexpr$or h .ohon e. difrcutt, c6ni.g iDpFv.s rhe p@isio. of th€ cults byimBing thc vitu load, bur this h.O.d i! iime @nlMi!& Iabonou& &d rh. lucBnle * low, hme the 6@h woi*6 hwc lo rely on ft€ p€rfotuc of plant marerialurdd th.n 6.U otrdirion arludons.

Althougb th€ imponm.e of nr dqclopmdt of CLCuD Bistmr @non vejdisha bs enphsiz€d by ney woll6 (Hwid ar at, l99l; MaiditL l99r: M.liL.,a1., 1995; Ali, t99; dd HusaiD .r zt, 2OO I ), ln. work don€ so fd is not comodable,and thb is pdhlpr du.10 l&k of irlmation @ ilrc g6eri6 of ClluD BisLfte incohor (Klh a dt, 2007). rd rh. pe@r investiSlrios, ile kiowl€ttg. of rhc clcuDger€tic! on rhi! sp*r he ben coI*rd Eing lle div6e cohon gmrl@ (T.bt. l).

Th. conp{ien of m@ vslG showcd rhal MAB_I l I had ln. tow6r diwfttin8 mdd Slsshow and fi€ld ondilionq jdicaring it,s bighly CLCuD 6ilr.oce

7a

.Q.rp.F.a ............... .... . ..,.. .. -q!e.u.tfB

(T.ble 4.1). in coftsr 10 NUCOTN-35B that w6 @ntrmed !o b. ridly s@eplibr..

R.ding srd ADulirg of 6tion vdidie bd.d on rhcn Bpons lo CL,CUD ainonlimed thcir higb cislm.c ed hi8h su$@tibitity of NrAB-l I I ed NICOIN 358,

r!+etiv.ly (Tdlle 4.3 ed Tallc 4.4)- tlc hjeb Bist&@ ofNlAB-t I I ro orlon tsf.ul viru di&e ba beo E?orrcd pdiosly by Allre €' at (2002.) dd Atnad a, ar(2003)_

ftc hodit€d djsle 6ting $al. us€d by Ahr.r €, zr (2m2) Drev.d 10 b.€filci@t in dettui tr8 vdiaiion in gmplM ssls.d ir rhis study, Th. dilrirctionbct*@ Urc 6isr{t ed sEcpribl. phtrt3 dong thc sir gdc6rio6 d.tu.ted w6 vdycltu Sad.l sturlie hrve p@iosty epon€d vdiarjon tor CLCID Gisr!@ bctwdO bobodeBe, G. hirsutun (Bi.\ 1975i dd Ebbcls, t916). C. htrrututu ed c.afio@ (Paq ad K\a2o05), goeijc dvds@ wiihin,idr!' l, ( }!l'ra 4r.2002 Ml Ailne et a1.. 2@2). In lhe pFmi 6tud, 40 @r!on edot tB qhibihrd

3i9i6@t vrialid b CrclD o.ble 4.1). Eetid FlDrrs by vdiou c.N[@ i..,siddiq 0970), Rehnd (2002), Ali 0999), Haida (2002), rqbat et ot. (z@3), r,tlja elat. Qo01') ttd Y\tw et ar. e01)7) pnvid.d Euffi.idt *jd4c thlr cr_ClD Bislsc in

Ih€ u@ of rhc di&s. deseni tc€hriqu. to sa 40 .oton goorypesprcvi{t d a b@ lilc for lh. a.lualion of sg.g.ring ald nm+grcgarirs @r!on

Smpla@ d.velop€d uMd rhc preht irv.rtigatiotu_ Th. qpoffs of thc sn mnof,g.r@lioB t, CLCUD w@ sihil& uldd ficld dd gL$ hoe conditioE (Tau€ 4.4.1),

whic[ @ bc aniburld !o similT l*€l of codeivc dvinmdLl @nditio6 such shigh huidiry. t€mp.6t@, $E dd vnd @rce (S-t2) ad whirelty lDflisia tabac,vctor, ft.1o8 6?oBibl. for (|l. rq4.id slred of ClluD s ,uga6cd by EioBG@bd (Giha ed Nou, l%9; Khe ar dl, 1998; Khe dd Kharl 2OOO: Altu r ?r

al, 2003; St|ttlm dd &dbi, 2OO4a @dtsg.rr. et 41., t93j.SiM 8@iJDi. 6!0ts ro $. di3.e undd gta$ housc ud 6cld cdditims

apPa&d io bc simih, only CLCuD dala coll@red udd rhe 6cld @ndjlions ws u.d 16

3tudy h. SaEric bai. of CLCID ard ilr .!si.tiu with oftd inDn d Dl&r

.QEP-lir-i ..,.. . ..,................ p!!s!r.t9r,

Th€ gendalio. mtu snaltsis mcilEd hs b€d us€d prcviouly ro study the

innaitara of 6is1rrc. b gBy laf spor (Co{€s, 1996; Cqr6 dd Wtrirc, l99E; edItompsr .r at, l9E7), and orhd dis.e! in com (Tond a, ar, 1993i Huch6 ard

H@td, l97l; &d Ctun &d H@td, I981a b) sd cisklce in corlon as.inri

b&rdi.l blighr (wdl&€ ed El-zik, 190). Dis.e s*aity Bring of th. six codon

go@lio$ tug€d 0.73 io 5.53 fo! the rvo p@1al cultivN md Ir poputation (FiSW

4,1)- CdolTE NIAB-I ll, diibir.d tignty Bisra.cc againsr @$or taf cd viudis6e, whilst the dod. cultiv&, NuCOIN-35D p6v.d to b. hiehly sus.ptibt.. fteitnd fd Da dis€& Fvgity vdu6 of ihc $CEerrjng populationsi BCN| ed F, ws idlhc diElion of highly r*i6rer p|6t @r..i.t (f{AE- I I I ed !r gd@tjon), sBg6tingfor lhe ddelopmsi ofplmi @tciial vilh cnh&cld Gi$arcc ro CLCuD coGidding drpdt h of vdiariotr qpBion in CtCuD by n ting deobB stetion fidr rh.sgEgating populalionr, IE .xpcsion for thi! plit€m of vriation in CrcuD inFercgating populaliotu wiu bc turoler sftngtn n d ion gd.-fo.g@c irt r:.rioshtlorheis a fuUy d*.loped by Bulconb. .l al. 1995 pldl viru3 parlosysr@s.

ft. edetic invGtigalios of rh. 6po@r of ln segregatjng ard non-

sgFg.lirS SaaalioB !o Inc CtauD i.dicared dst rh. fou-ptu€t€r model m, Idl,lhl, ed Ljl pbvid€d 0F b6i lil of th€ obsqryed ro thc *p€cl.d sdturion mes (Tlble4.6)- D@ io de gqrd tuAnitudc of rhc [d] @DpoMr the inhdjir@ of CLCID i, tbeplbl matdial app.aEd ro h. influo..d by th€ 8o6 &tin8 addilivcl, sinild tofindings of Ombl., { 1962a b)i uoe &d w![er ( 1968). rh6c wortcB reDorr.<r rhe

imponft. ofadditiv. 86!ric vdialion in @droling F.its $al @ ls coftpld ir rhcn

'nhdila!.c, Bsed upoo tn€ g€nc acrion conholline CLCuD, it s@ thr rcsisla&c for

CrcuD in thc plar Eddiat €lmiled n@ may bc inprevc<t by tuIir8 &letid of the

dsirEd planG in laid gen€rarioN, s had bM done by ciha ed Nou {1969).

ln prqs.nr sludi6, ft€ adlitive gdctic vdiec ws sigtrific&dy hiahs llta itEoviremqLl vdiec. for 1h€ CLuD r@lion (T.btc 4?) in contdt wirh Epons ofSiddiq (19?0), Rclmd (2002). Ati 099), ed Haid{ (2002), who obw€d ihar

CLuD ws nainly @d6 rhc 6eot of singl. domiut mjor go. wit! iNolvd@r ofmino6 (nodili6) 9o6, ft6c @ntrsi rcsutt unda Ue pGdt sludj€6 may be du. roinc llsn tbr onon poputa,rioB ls€d udd th. plt$l irvdrigrtiotr v@ difffit

80

.q!sp.F!5

filn th. popllatioG u..d by Siddiq (1970), Rehrtu (2002), Ali {1999), .Dd Hridtr(2002) i! ihcl previou dtodies fm tbe gdclic sMi.r of CLauD Eislarce. Ra rhawa

(199) foud two donindt e.c ro bc iorctved in Cr-CuD Bis!.nce. while d@licdedomiMr €pirtdis hd b.a Eponcd b control io CtruD @i.t.ncc by hbal .r al(2001). on 6. oid.. hsn4 Atruj.

', ar. e00?) ..pon d invorv@@t ofth s@ wirh

duplicare tlohi@1, domirel iliibitory ed duplicatc @6siv. epistasis and th@ ga6wi& oiplicArc domi@r @isrsis irwlv.d itr CLCuD Gisl{E..

Thc pE@t dam snow 0E pt€gc. of rc8arivc donildce [h] a w.[ s the

'Nolvfror of n g.iivc addiriv. x doDimce tjl int@tio6 foi tre CLCr|D tnir. Itr€n€Salive thl wlue irdieEd lt t domiec w6 in th. dnatiotr of Bista@, whilen gativ€ Lil val@ ddoie! rhat its.ff@1ws neslisiblc in Educirs the additiy€ vdiece.Thc highq v.he of addirive wiee 6dhd subsladd.d thc inc6itr8 tredisibjtityof 0E go.s with ircrcrs€d reist m. against CrcuD in fie succedins gfld.rions(Tsbl.4.7). Hid addirivc vdi&e |str€d in hish 6t'dd6 of |t@v s.@ hai|dbjtitydd qp@t€d 86c1ic adv.N for rhe CLCuD @ist.G (T!bt€ 4_9), lueg6ring !,rtpFgmy sl@t'd would b€ etut ir idorifyine plelt wirh bc@sed CrcrD r€sisimeudd ni8h it|@lu lord s $ga6r.d by sinsh md Ndyr@ (2ooo). N.r!tiv.h.t bli€ €ff@t for CLCuD prcvid.d eviddce oI th. p6oc. of addidv. g@6 b.insdiretioEl rowlrds lowa pffi1 (I.bL 48).

Cdqltion ntus aElyri! for l€ll e. ed fibd lagth r€vealed $. adequ&y offivc p!@ad nod€l tD, tdl, til, !l sld [tl, bdicaliry rhat bolh traits wh @trtro .d byu€ gm6 &ting a.ldiliv.ly &d compticsl.d by !.gativ€ dd pcitiye jrr.@tion €fidlsof .dditiv€ x additiv€ [i], neearivc additiv. x doni@. Ljl od posnjv. donimce xdonin.@ [] inr@iod. Th. sigdfcdr rdditivc g@cti. efiec to.lof rP dd fibGlo8th suegBted that si6pl. *laiion @uld be mor ellaliv€ in jmprcvirS the

ch!ruld in lr,Ia g.!@rioE (Sing! ald Ndluo, 2OOO). Sinit r id!dir!@ qndfor fibd lddh h6 bdn rcpod.d by nmmus voft6 (Shoky .r ,t, 1986; Vrdalkd er

aL 1984; Li! and aa, 1988; Nldrllie .!d RareM, l99O ar|d 1994 W&g ed pr4l9li dd Md-Zho .r al, 200j), Itc n gativ. .pislltic comFon@t lugeerr rhal

fix.tioD of lddirivc x ldditiv. 966 nay r.due O. Ieaf @. in taid g€l@iiorrsuSg6ting thc iddrifclrid of individud plra$ wirh l[. &3iE<t taf @ woutd be

8l

.qrP!9.i....... Drsc*im

advetrgou in dly gderarions, tn @iru! th. posnivc addirivc r addiliv. [i]nle@lr@ s!98*rs rh.1 rh. fxation of addnive all.lcs for fiba l€neth would bc pcsibleiD rhe later segrgaling gcnaatioB, Ibe signifcdr I for hon w.ight luSSest th. n€€d formi.ing tuc€€dine gdcr.tioB in oda 10 ddmir€ rh. b6t gderic nod€l (M.tha and

Jint5, 1982)-

Fivcptu€ra ooder, n, [d], Ihl, [i] ad til app€&d ro be ad.qur. for p.tiolclogth, rubcr of boll6 ed *d .oton yi.ld (Ta!I. 4,6)_ Th. gdetic anal'sis of lh. dar.for t!4 itE ch@i.F dotcd rhar both additive ad domi@c. g@ cfrelscontou€d tho. t6irs. h additior, the sigDifi.el losirive .ddiiive ! addiiiv€ inr,&tionlil for ib* thE tr.ir. 6ugg6t 1btl fift,rjon of f.rclEbl. lddiriv. .I.16 would h.p6sibl. in lhc lala 6.gtu8aring s@rions, 6 sugg6t€d by SinSh {d Ner!fu(2000), IlE irrclvddl of boIh thc lddirjvc dd dodil|.e gcn€ etreiq ed ddirive iaddiliv. [i] inra&ri@ .pistatic .f..1in conr.olirt itr€ i.ncntafte of nubr ofbolsdd sc.d @fton yi€ld e in grcar @.&miiy Mrh fi. stldies of Wdg sit p& (tgt),D6bp.nd€ ad Baig (2003), M.n 4 dt. A@32), M.<:rj;ly .t at. (2004.), M.r.dnn(2005), ud M.|zn6 et at. t2oos\.

'Iho d.r. @ synpodiat tqgrh, trbc fild*s, flbo tuluity, fibd s.rgL andnon-s'pifi.e. xa vdu. susgq! the a<rcqucy of fiverEmdq mod.r n, tdl, Ibl, tiled (U. Gdclic ef6b 6tin,r€d for rhe trajrs @eted rhlr 0E addjrivc ard non-addiiiv. ed$ @ntolled fibd lFogltr hoNder rh. ctI@ls ofnon-additive gcnc! w@noE imponq! Stnpodial lcngtb, 6lE fnfls ,rd 6ba ll.ruity w@ l&g.lyinnuftcd by th. gd€! &ting .ddiriy€ly. ,Ite low lddilivc gdetic .ffetl contoltingrtDpodirl lagtL fbd lind*, fiba rutuity ard tiba srEgrh sugget th, slerioo i!dry sd€ntios could bc mor. .Iralivc. Il)c oppolitc sisn, of [hl ed ttl for dl ih6efou i..ils iqlicaled th. iMtvcnal of duplicaL @ist sir. Ihe n.gativc cpirlariconpon€trls [i] for sFpodial tdSrL tibd fM€s &d fbq manuny @.al.d 6rrfxatior of addnive x addiiive nAy rcduc. the srmpodiat togi!, nicrenliF vatu. dd@tujty (lowad. 6G fibq wilh d6iEbl. fihs matuity v?rlG) of fb6 in lard8oaafto6. Thcrefor€, scletion of individusl plmr! with thr€e chdrctd would b.

8)

Amng rhe dig.nic @isEtic cf]e! rhc s'gnifitut pcitive addiriv. : additive tilirt @tion elfeG for 6bq shd$h sugs€rr rnar tn fiiadon of favoEbt. indftivc dtet6would be po$ible tu th. laid egr.galirg 8@hrioG. Thc godic mrrot of 6btrstagth h rh. pEor wolk b in ag€mar wirh r[. tudings of TlBu 0982), a

Mvre d al. (2004), who rcpoded tt t boih addirivc and donimc€ ef@k wftioporfit i! thc irldirture ofrt€ lrir !n addirid, thc .rudi6 ofMc Ctry d at, (2@4^J

md J&k.r at (2005) rcved€d.ddiiv€ x addiiive (il ilr@ftr w@ inpoiant in the

irhdine! of 6h€ srdgrh. R6utt5 obioin€d tlD it pl!@i sru{iy shoe l[.mpone* of sdditivc gaca, ed $nc jnvotvcmdt of @isi6i! al doni.s.c x

doni@@ l@i in @ntbtlina fiba find6 s[t tiba naruity. 116. finding6 e inr8rm6t with pEviou rcports (ar4 a at,,1914i Lin d\d zie,l98Ei dd Nad@ieard Rd8asy, 1990 dd 192).

Ad€q@y of fiv.-p.mdd hodel, n, thl, 14, Ljl &d ltl show.n flc .lsfte ofatlditivc gactic elfel3 in rh. conr&l of bou sjza Thc msgdiude of [t] w6 grclr.r rlotL msiitud* of Ul, til and tbl. rhe oppcil sign! of [h] &d lll for roll sia deindicaLd the crpGsion of duplic.r. episcjis, $ggcrring conpl.x inldn&.c, 0r.rcfoEelerioD of d.sinbte boll size @uld b. .@Dplilhcd in @ry ea€r.r,ons,

.r.hc D!!grfindinss @ in conhey ro th€ lrudi* of Chsthm er .t (2003) pho foud bol)t a<bilivedd dohitll@ g6€tj. cfldr to h. inpor&r. Hoecvd, rhc pEgr d.rr is in a8tuarvith Rddhawa e, al 0986) andings.

Ihe fou-p@.r.r nod.t, q [d], til and til ws & ad.quale ft fo. sin rh out

FMnrae. d.o. Th€ ga€ric e.ltsis show€d rh{ the addirive ed.s pr.tohiettyetr4t d p€rchi gh ru our. posiriv. addiiiv. x .dditive [i] cllcrs wG idolv.d in th€inlsit-'n@ of p.rcat 8ji nn o!! indic.ting thal nxatio! of flyoEbt. lddirivc al€lesDay b. posibl. in 0! lat€r FgEgalirg g6drjou, s sugAelcd by Singh ardNeyrlm (2000)- Thc ptl.4c of .ddiriv. dd digdic .pirtlrjc efet3 for p@.nr ginnrn out hq ben sbra iat d by the work of Sirah ed Singh (t9Sl), Kasm er at(1984), CherhaD a "l

(2003), Mqr et at. l2@3d dn M{$ty a ot- (2@4 ,t.ft€ finins of rhc sa€tic hodet using gmmiion vdi&c6, addiiive [D] &d

dlinmoi.l {El @m!o@rs sem€d b hc luficidr i, qpt inins lhc vdjatio! i! mslofih. !!it5 exhin€d, Thc aig|ificst dd tarSd D @hpon ni iddicaled rhe dispdion of

Ch.Drs s Di.csim

E.l

.qa.,p!r.r5 ..,....-. ,................. ........plr9t!!!-

positiv. ard n€g.tivc all.Ld in rh. pamis, A darisfa.iory flt of rh. model incorpoEtirs

otrly D &d E in 1h. a6@1ion edil@ araltsis slg8Eds rlE I,qae of lddirivc g@dj.veialion for all th. Fail.. Th.e ob6aatio6 indicaL liial @nti!rcG *ldrio. ce be

uFd to idmnfy desinble ple1r. A eodal .d€qud€ firlcas of th. two-psmetd hod€l(D ard E) b ln. gmtiotr wi.lG sugg.'t' thar rdditiv. vdidc .ppar.d io &@utfor lhc 8reatq p@ponio ofrh. s.ndic vdi&e for sll th. lraiB .rudicd.

In ihe precnt study, ihc efnatca of n@w seM hditabiliry for au ihc 13 rnir!e pMled in T$1. 4,9. Higb nmw &@ [dirrbiliry 6.s) 6dnar6 fq gi! nmoul bou sia, Laf aE , ot!o. lcaf cul di@c, 6bq lino6s, 6b.r maruiw and sed6io! yield reflecr.d high g6.1ic adv&cc. Ile iiodqrre hdit bility qriln|1. forNabd of hols pd pldr suggGt d thd gr.dDaly iopovddr in rhis tr it @tE be

oadc ltrmuSn rbw pDgEs. t1i hign |rry ffie hdjLbitiry qrimat6 i.e. .xcc€dine

mity, for slmpodj.l ldglb, bol r€igh!, 6bq logtb &d fiba rrugrh. nighr hrve

@un d d@ to vdiou &atoB $ch s; p6. @i4. 6rim.t€s, pr6.E of mn{I.licule@lion, @pling mr, dd prcbabt. prc!€@ of non-gaetic aoe6 of vdiario6(Coaie .rd \lhitc, 198).

Th. 6tidc. of t':s for Ftotc ta€tb js ood@re IX. highd 6limr6 ofhdiiability for ihc ch@16 .xmiled hft nigh b€ s 6u[ of gEara edetic vsrilrionof g@6 Mhr€d with cr,cuD, s *eu]d.d by samg! e, zr (199)_ BE&hv e.rIi.dwick (1989) .rgued r!.r uhou vdiatid in t@l61cd halqiat .orld be

u@va.d whs noddr€ !t .s h appli.d wbich could po$ibly increse hditrbility.lal@ld ed M&t!, (1996) $gg6r!d rhal hainbility 6tiD.tca @ 5ubjer tocsiddble aviremor.l v&idio4 .ad r!@foE caurio! is EquiEd io dala

int€lprcialion sd ue, Noncth.le$, thc high of hditrbitily .stimai* und€r thc nigh

CLuD ineulu lord sa ro b. rlirhlc drr. llar @dd b! ofvatu€ in br€€din8.

Hchric.ffors w@ vciy widdr in sI ihe rni6 cv.lur.d. N.g.tiv. h.t.cisfo. CLCID, ilbEding d.prcsion ed ,otcnce redo previded €vid€ncc of additivc so6@ntllling thc CLCuD Gisartua Th* 6ults iugaBr rhar CI-CID 6in4. irCostnirn iimttn L,,n ! b. imptuvd uing ln. ptlst phrl n rdial s h4 bend@c by Giha dd Nou (1969) b Sudd. TIE rcgaliv. heifrlic .If!* for CrcuDlslrtu ds pDvidc widc@ of 8a6 rdds h l[. diEtion of tn. hidy ci,i.!t

E4

.q!'r.p!9..s_................ ........ .... ......p!{9!l!d9[

pami. fte rcsativc inbFcdiDe d.pBeion fo. CrauD r.si!r!e. als suggcrrqd ihe

plEace of treg6sivc s.grsets i! F, !d b&tclN sEcrdioB.Thc @Um.. of bb! Drgnilud. of lttrid vigor fo! lef a@ dd trMbc of

bolh ud6 feld @rditioft night be du. ro g.n6 witb pteiotropic effsts, Eigh po!.nce

valu6 for leaf e4 p.tiole ldeth, rMbd of bons, boll w€ight dd bolt sjze indicai.d

thc Flpondm@ of noHdditive a@ .flsrd rrd in $D. *6, th€ involvdqr ofdupliorc €pistatic !flet3 i... dooi|),e r doDi@ ilGdtion. Negrtiv. [cr@tic.f6t @oqdi.d by n.gdive ilhleding d.llsiu fo. lynpodi.l lagln pDvid.d

€vida.c ofnldiriv. goc cf€cts ontrolting lynpodia.l l68lh, sugg6ring tn€ pdribilityof sloling longs srmpodids witn Sood bollr in sG*ding 8@r.rio6.

Cml,rtio! @fficiat! fo. veioB !!ter.d wi.r6 .mng th€ sn-BcrdtionEaa 3[ow thlr tl!@ *6 g@d assidid emg lriala $gg6ting thrr ildicr$leriotr.d b. us.d ro in!rcve &y of rh. rr.its, the gdor,lic cpoIB ro ClluDsho*.d negrtiv€ coEGlation cGficimis wiih nmba ofbolk, bui had sisnificst edporitivc ssiatio. with fba 6no6! erd gi! rM our, Thc disnfiool positivc

al*ilion of CI-CuD wirh ni@niE v.lrc indiore &.t CICuD irfeted ptdL hrd

Pd qudity fibd. It. ne.tive @reLrio. of CLCID rilb rub6 ofboUs signifd th.dv@ .I&.8 of CLCuD @ ptdt so*1!, lad ullsEr€ly on nMbs of botb 6rta rh.prevdling CrruD inocoulm load. Lcaf e! dlibited highty 6ignincdl posirivc

$@i.titu wil[ p.tiol. ldgth ed nmbd of bolts, ed si8lificet pGitivc 6wi.!ioBwilh bol weighl ed lad @notr yicld s!8g61irg 6.r *lerior ofpL s will Ecdiwlqf .@ eith d*inbl. pctiole lagrh is p@ibl€_ nre Dosiriv. .!si.ri6 bdwM t4f@ dd p.!iol. lotil n.y bc of val@ !o irr. bEcd6 ir natjng pbtrt slerioo with

high nmbd of boUr at U€ lowd planr csopy. I{s mly f&ilibL ro lMiniz. $l&inr.Erption leading ro inc@ed e€d @no. yicld.

Th. Fcitivc aei.tior bet*6 lef .g ad s..d @rion yi.td ob@.d in rhis

srudy i5 in @dlict with ln. e.lid findings of Dhoptc &d Jd.dr 0989), wlo .@oncd

n 8aiiv. @@lation bcfw.a l.af Na dd hd.sr idq &d sed @non yi€ld. tt.strong positive a@htion betwd ield sd l€af @ may b. due to longe p€tiot. ihst

ould havc f&ilirar.<l lEinM had.bL bolh noD t!. Iowd h.lf of the @!o lltutrl naluity, Ih* aulrs sgg.!r!d tul taf .ie @utd be dploirr&d dity in tutua

85

S&p.le!! ........ .. .......D,.9y{,".1

bEeding pregm€s s srgSE$lcd by Ji.ag d dt (2mo). Th. hish siif@r posiriv.

s@ialiotr b.tw.d petiole l6grh ed nunbo of bolr and ,ignincd posirive

.lsiar,o6 of pctiole len8h wirl hoI ecigh ed ,..d @ilon yiet4 tu thd tuppo,& 0l!tadEcs nay b. mad. by qploilinS thc Dot@iiat ofptmt daterist sudicd undq thepG.rl im6tig.tioE SiDildl, rlE sigifitur politiw asocirrio, ofbon wcieLt ritbnuhd of bollr sggBt d ilr€ po$ibitjty of @tring inprov@oi in rnce traits irtw..ding g@drtiotr. Thc po6irirc .!soci.li@ obrdin d for 3..<t @iio! i€ld Mthomb.r of bolh ed boll weieht j5 simil& b d. findings r@ncd by vdioB wor|ts(7ytsi, 1987;76nt a at., t992; srd Ty.gi, l99a a, b),

Ia mcluioA rhc Eulb indicaled lllt th@ is sigdfi.or 86ai. veiobitity in$. oaLricl studicd Tn @tron culriw NIA&tIl, Fr atd BCM lppdfiioB €xlibiicdr.!klae. b thc CrcuD udd borh scb of ovibmenls. Cultiv& NuCOTN-35Blppdld to be hiSl y swcptibtc wlliln Fr m<t BcNlFpul.rio6 snry.d s.g@gation for@tton l€af cul viru dis.sc, mc CrrlD show.d ft€ n.gslile afl61s on tal ar.a,nbb. of bollr {d fb€r fM.sg lte ptgat ,rsutr! sggdld rh.t dis.e 6isr!!c.of pl&i mardist sr.ic{y bas.d or iheir ratr8s 10 CI_CuD, itei ed€tic bsis ddeiaton wih olld r.its @utd b. u!.d 6 *l*rio, dirdir for UE it rclopl@r ofh'Cy di!.e rctisLd conon a6ot)te. How.vd, in ordd ro $b6i6tiale rhe pEarGrtrs nlth€r ir!€dgrios @ r.qui..d befoF maLjng erdon ,rcgrh@. .ft.g.ndi. idfonalion daivcd non th4 siudi6 tunlld s!€gcrt that rhc rraii! .ondirion.dby tle 806 wirh addiliv. p6pqti6 woutd tE h.tptut for iopoving thc Er&.iald.veloted by ft$ing dive pa@b.

86

Sumrmv

AllouC Cl-C'rD eu ida i6cd i! 1967 i! Mutt ! otor!.i! sd Ari, l9?5), nwr @t ! pFtLd unril 1989 &d t990 dE !o it dinor iDlqtoe Howd, durEalha yrs diqlc iEid.occ isE&d @[i<la|bly cuiry tosr.. @tiig 22.3 ro 58.5

% i! ldE ffi of hnj.b (A@ R?, CCRL l99G9l; Mult ! (prlida). Di.....ecity daadd on culriv..r sFsn rie of idddor .!d th. growirs ..odili@(Hu$.i!dar.,l99l).

I! vicw of @tbn FD.lErioo ptDbtd in pd.id.a th. pc.6r @:rL wdt qscai.d o'n !o idcorify rtE .ddiriond bor... of Bi.rtD &rd lo iddtigd! r[G

iohdit|n c of CtfrtD @uing i! Pitido_ ID &. Fsar iftrdisirio!' di.c& dinSl\tclniqE b.r.d on thc 0{ .cdc, dcv.hpcd by Alhr.r a zr. e0ol) dd AE r rd Kb6(20@) or! u!.d io 6du.L rbc t {o9Dic n pdr€ !o Ct CrrV di!.G..

Prlviou srt itoG d $. irlditdEc of rlir vir.l dig& i! sc. y, .trd wblldo€ ci.t i! liialle rtow rhrl CtauD b coorDlld by. singtc ddEi!.d B.d. (Ali,

199; Rdb!4 20O2i lDd Hridq, 2oo2). o! rL 6rrny! p\odbu (199) lnd lqb.l aar (200!) rlpond 0d CrcrtD w .ontlollcd by ls doDinrlr gaE Dd dupti€r!donimt .pidr6ir, Elpetircly. Tt. 6!di!8t ofK!& ., a/. (26,07) Bat d lhrr ddiriE8@ crc.r! coutriblr.d nd! b 6id.ee

b rh. Pt!s.! rtudy. .ir @dor g.iddo$ ditrqld in th.ir r.{@d lo @[.dlef curl vio! di!a&. TIG coion cdrivr NIAB-lll, Fr &d BCrr popll]|li@Flpdd.d frrcnbly b ibc cotro! lafcurt viro did$ uid.i bolb 8Ls hd4 .!d 6ctd

coodiliooa vhild tb. culdE NuCOTN-35B @ai.d ro bc biCy e!..?libt..,nc F,dd BCft popoldioo! m&d in rbcn Erpo.,!G. io coloo lefclrl viro dis*e

Cdic e.lyti. i4lic.Ld thr ddiriE 8@!ric ctfd|! s@ iolub in rt.inlGildcc ofc<,ttd laf cud viN dira!., egSgtinS 6c cbrEla !o bc FlyS.dic jrEte Th. r.covqy of CI.CID sid.@ pt6a i! D-tcBir€ t6q!i@ s8S!.r.dtrlr tn. cl|lat r it bisNy h.ric|!l.. TL p(etr bfotuni@ d Slncic @b.!id ofiildiLe ofCtCoD, disSr!€ wirh U. r.ponr of Sitdiq (19?o), R.t||u (2002), Ati

8?

.91.{pt9r.6,... .....- ....... .,s.gt9!3.r.y-

099), ard Haidd (2002) who foud CLCUD b b. @nroted by a sinsre Er&rdod'@i edc with ninoE gqca, IE Eor findings undd rh. prcsot studi., ,greewii! u€ 6rdi!8! of R€hl@ ,, at. (2005), I<ne .! dt (2007) ard A!uj! ar dr eooT) whoEponed th.l rddirive 8oG etrets @nribut€d tI@ ro ctluD cists!@. Thcinrclvd@t of ninor sa.s n.y caue veiariG in discaj. dpE*ion (Huichinson mdKnidt, 1950), which ce c.ue sscAtiblc !&ieri€i with f6lq growih rarB ro c&!p€6'ly alt&ting of CrcuD rh& do slow gbping 8@tD6 (AI, lg?} Ottd wo*6fohd th. involv@ml of rwo do-i"or S€n* (RddtNs" 1999) nt two dominulduplicale gd€s (tqbal er ar, 2003).

Adttiiivc goetic vdislion ts at$ bq suge6Ld to b. a eFsrcr irtrporrrrc. in|nic whi.h d l.$ conpld in thei. inhan rce (Ctrblq l%2!, b; Lieg mit Walla,1968). B6ed on the inhsir.ace parm of CLCuD, Esisrrncc in cotlon D.y b€ inc@ed0'Dugn dBightfo dd pbDi &t*rio!, Thc hj8! n g.tivc doni@t go.rjc cfids 4wcll 4 rh€ invol@ent of @saiiw [i] cdpoar irdi..r.d rh.l doninee ws in thcdtr ction of Gisrdr peor. How€vd! snalt negariv€ vslues of 0l &d he&oric efiei

'ndical.d thar doniE@ would not Educ. fic nsg tud. of adtliriv. veiaG, ad rhN

to inclle d6 posible rdsisid of CtluD Gisrdc. in rhe tucc.dirg 8@d.riom(Walla.e dd El-zik, l99o).

fte hieh $tioate of n,lN-sde hqilability e@iated willl higbd gacricadv.&. for ct€lD |@rio. uldd hie! f.ld inoutm told seos !o b. €mm8irg rotnc breedd for mating &l*ri@ oflhe pLtrts Mth €nis.cd r6ista.. io Ct_CuD, Ih..fi6tilo6s of this siDr.sl hld b€d @phrsized by bight (1943), sd Cih! ald Nou(1969) in Sudrq vn@ t}e suc *leiid pd.dw ws @trriDuuty uscd in *@h ofhighly r6isknr 1itr6. Cu@r Eslrs irdicai. lhrr pldt marcriat u.d in rhe snr(ly may beof vrlue in d.v.lopilg CrcuD r$isrsrr cultivs. Thc high l)mw_eM h€rirabilirydrtur. for agEnmic &d fiba quatity mjrs, $eg6r.d th.t ir i. posjbt. ro idcnrify6d!m pleB wid d6i6ble eonomic chmrE nom F, &d bekcNs prcgai.s,

Laf e. ed fbd lqgth oppcared to be Ddq the i,fl@cc of 0F 8q€s @lingaddniv.ty alons wii! dia@ic irr.@rid!, n.sarivc e.t posilivc [il, n.cative til edporitiv€ Ul compondr eff6i. Ihc p6@. of signif@r additiv€ gddic co-IF"qt forbaf aru Ev.alcd fiat sel@tion in llt€r BodrtioN could bc nore €ff{rive in in revinc

8E

9!fg+{tJ

th€ cheEler. Thc n 8.1iv€ epislaic componat [i] eugg$ls lhat fix$i@ of additiv. x

addilivc .fiets nay red@ lqf @ in Lter gatdrti@. rrw addiriv. gdcric ef*rs for

6bd ldgth indic.!.d 0[l *letior of ildividu.l pldts qii! befter nbd hSrb would b.@ in dly 6trg6. Th@for) dly 8@ati@'. &letion of pldrs *iln lowa lefa6 @uld bc bcn ici.l. Th. p6itiv. [i] .oEpon6i sg861cd rt l fixrrion of artditiw

all€16 fd fb.r laath in later egreSrriog 86cr.ri@ My b. posiblc G srggBted by

singh &d N$rym (2000).

The 86aic salJ6is of p€tiolc lcngitl nmbd of bolls dd $.d cotion i.ldHeld rhalrh6. tnib e lrdd the @nrol of &lditive dd doDimc. goe efr€txwih .r.nin,rc goc cfficts .pp4i!)g to bc n@ i4{,r.,1. ft. ptg4. of sigpifie4posiliv. addiliv. r ddiiiv. til in&tutioE @!oMr for dl rb* rhc t6il5 sgg6:r€d

lhat fixation of favoRue .dditive .ll€ls fd.ll rh.e Fail3 bav bc @sibt. in the |rt r*gregaljng g€n€di@s (Sirgh dd Nmyr@, 2000).

Th. g.nctic .'lots for sFpodid lqnglh, fiba ftno6 dd 6ba Banliry

ry€d€d to b! uadd tbe .onrDl of additivc 8de efers wirh pFdomirdl infludce ofd-Ein'i@ rac cffets. Ttc appGiie sig of lhl rld lll for lh. fou tni|l irdiqi. rnc

.xpBio of dutlicd. .pistair Ir riifmr pGirivc addilivc x rdditiv. .pisrrsb [i]coDFMt fof libd rE€trgtn sg86t€d tld firtrio of favoEbl. .dditiE atlel* fd bod

clrdEct€B would b. posible in lat . *8r€gaib8 gm.tltioB.

Ite two pamdd mod€l of D dd E lined b lh€ go@dor vdiec$ sugg€sted

ilal arlditiv€ vari&. &@dted fq ! ldtd pbportid of loial gsteric vdime for att

llc tai& sudicd. 'ftc siifor &d ldgd [Di @EFrar Hab ttrlr bo(h posiriv€ and

n garive allclq w@ rc1 eiatcd with @ p.6r This b.y b. dE ro oe p!6e@ oftbl dd @ft@. of high nagnitdc of hdmd. in Fr hylrids p6nicll&ly wirh !@b6

Si9ilicdt positive a@iltione b.twca Ftiol. l@8lh dd lcaf e4 nmbs ofbolls wiih l.d &t , rMbq of bole with pcnob l@gl\ tbc fnd* with 6bE

D.nuity, !.cd @non i.ld with !€{iolc l6grlr nuba olbof with boll weight dy be

du. to lh. lilt gc b.rwq ga6 @tolling rlE lllir& tlryffi th. plE€e of CrcriDaplE4 lo ltFw. rcgtivc lsilrio! with laf .M ad nmtd of bolb. Sibildl, th.

E9

qr.,.P!!.{.6. ...,99_--'"t

signilicrn positive alsoci.rion of CtauD *irh nimdirc v.lu€ indicatd fi.r CLCuD

'nfalcd pl&ti h.v. poor qlatio/ fibd.

Th. pre@t 6ultl dugg6r€d rlEr Sderjc vdiabiliiy for CtalD &d othachqr6 dG qists in rhc plaDr rn di.t EdcEted by crwing tso dirtE cultivGNIAB-I I I ed NuCOTN.35B. ft. krcwldgc lbour l[. gmcric b6is of CrciD ddp6iliv. a!@ialio$ betw€d d6nabb chucid could b. ustul for Eaking &leior oIplea @Dbined wirh d6inue 6bd tr.its, The infomatior r.poned h@in uy be

dht 86E to @tlor bF.dc6 for @lrinucd iDpovmdr of clcuD 6is1aE. in

Gattpiun hitu L, nnAd lcsl oviomarat oldirioN,

CO N f RI B A TION TO WA RDS EXB T NA KNO W L E D G ESiee l[€ pEviou irfoD,li@ o rhc 6isle@ ofvdi.bility in CLCID in @tion

is rol drcnsire. The pc.nr mrt ..po& ddlit d b&mation on rhe r€.poM ofdifi.mt v&idi6 ro clcuD. Ir* g€doit?ic rcspoNd io clcuD mtcd h@in @signinca suegesiiig $e po$ibiliiy oI uitrg rhe pr€ffii g.mpl@ to dw.top Ct CuD

Eisi.rr mlaial thrcud brccdins.

Th. p@iou rrowl.dge of gdcric @nrot of CtluD k vay s@ry rnd wtltd(B dilt in litl?rw h.v. b€o .ldivqd $ing Fr &d Fr poputrrioG onty, Ili. p@ntilfomation on 8€o.tic balis of CLCuD of g@orrpic Bpon$ w6 dqclop€d by

evduling six g6fr1iou Pr, P,, rr, t,, BCM ard BCN| rhnuStr 8@.ration htuddFir. 'nrftf@ lh. p!B6t bowLdSc of gderic bs6i5 of CrfrD js D@advrr.g@E alkl etul b @nd bF.d@ lor llt€ e 6Md1of craiD cisr.nc. ir.otion nat rial thrcud brcldira ed sel€ciion.

PBiou idom.tion otr ad.dc bchavior of petiot. ldgih in prad poduclion

dG iol qin in $c liiddue, ard r[@&F th. ilhdillE pafim inv6riAakd n@infd lhb chmtd is nd on.. The sa.tic infom.tion dcvcto!.d hft, in colttlllins this

bait, will bc ofprime vdu. fn .otton b@d6 whil. Daking el€clior of desidble Dldtsmolg egrcgating populdios.

90

.-CIPF-.6-..-..-....-..........,... .,....,............!!t'*@.r.1

SUCABS''ON$ FOR FAIABE XESEZIRCII

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