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INHERITANCE O F STEM PIGMENTATION AND FLOWER COLOUR IN CHICKPEA (Cicer ~riefinunt L.)
KlRAN KUMAR V.S.S I3.Sc.(Ag.)
MASTER OF SCIENCE
I ) ] l 'AK l !d l N l 0 1 01 N l l l ( \ ~ Z ~ l ) l ' l A N 1 l3l<l I l ) l N l ~ ~ l l I (k l '1 4 IjIKl I I ) lN<, I I N I I ( 0 1 I I(II 0 1 ,Z(rIIICI I I I IR I IKAII N I )K IZ~~>ZI IAK I( ~ J l \ , \ l AC I l A K Y A N (I R:\N(r,Z A(rll lC I I l l I I < A l I ' N I V I !<\I I Y I'A I A N ( Ill I < ( ' I I Y I I I I IAHAI ) ill11 llill A\I)III<IZ IPR,ZI)I \I1 INI) IA
OCTOBER 2001
CERTIFICATE
hlr. kllt:\N k l hl.\l( \'.4.S.. II,I\ \ i ~ ~ ~ \ l i ~ ~ ~ o ~ ~ l \ I)IO\CLIIIS~ IIIC LIIIII~C (11 IC\C,IILII
,111tl 111~11 ~IIC ~IIc\ I \ LII~I~IC(I " INIIFl<l l:\N(.I: OF S I Fhl I'l(;hlliS I ,Al ION ANI)
l~l.O!\l~l< C01,Ol U IN (' I l l( kI1l:,\ ( ( ' ~ [ [ v ~ I ~ I L ~ ~ I I I I I I I I I- ,) " \III>II~IIICCI 15 IIIC
rc \ l~ l t o i i111f111,il ~C\C,IILII i i ~ ~ r h ,111tl I\ o l \II~I~LICIIII\ IIILJI \ ~ d ~ l d d r d 10 \~,IIIJIII 11\
~plC\Cll~.IIIOII h l 1\11' C\,llllllld~lOll 1 ,1\\0 LCIIII\ 1I1,ll 111~ IIIc\I\ 01 1),111 IIICICOI 11,1\ 1101
I~CCII I)~L'\ ' IoII\~\ \~I~IIIIIIIL~ I)\ 111111 IIII L / I c ( I c ~ I c c ~ I ,III\ IIIII\CI\II\
I l i i \ I \ lo cerlll! 111.11 ll lc l l ic\ i \ c t i l~ l lc t l " IN I IE l< l l ' . \N ( ' l I 01: S T E \ l l ' l ( ~ 3 1 E Y ~ : V ~ l O X A N D FI,O\VEl< ( 'OL.Ol~l< I N ( ' l l l ( ' l i l ' l ~ . \ ( ( ~ c c r ~111c111111111 1 ) '' i \ \III~II~IIICII In ~ , l r l i , l l i ~ ~ l l i l ~ i i c ~ ~ l 0 1 l l ic ~ C ~ I I ~ ~ ~ I I I C I I I \ It11 l l ~ e (lcprcc 01' \l:\S'l l<l< OF' SClF,X('I< 1% ~ t ~ ~ l < l C l ~ l . ' ~ l ' l < F . 111 1I1e lcl1~11!:1 V ( I
1<:11ig;1 \ ~ ~ I L L I ~ ~ I I I ~ ~ ~ I ~ l~ \c l . r i t ! . tl!ilcr;~li;td I \ 11 rccoril 0 1 IIIC Iho~~:tli(lc re\e;lrcI~ \\orl\ i.,ci-ricil oln Ih! h l r . l i lR : iY K l l \ l A I < \'.S.S 1111(ler IUI~ ~IIIII;IIIC~ 1 ~ ~ ~ ( 1 \11per\i\l1111 I IIC \11Iijcct o l ' t l ~c I~ IC \ I \ II,I\ heel1 ;lpyl.rr\cil 11) l l ic ,ludc~lt'x :II~\ i\or! c r ~ t i i ~ l l i ~ ~ c c .
TABLE OF CONTENTS
-. .
S. No. Titlc .
2. REVIEW OF IJTERATURE
3. MA'I'I7RIA1,S AND MI"1.I IODS
6. SUMMARY
7. 1,lTERATURE CITED
LlST OF THE PLATES
Plate No. Ti t le I'agc No. . - - -. - - -
I . Parents s h w i n g ttcm piglnentation ICC5716and ICC'17101 34
2 . Parcntr rIm\\ing flower colo~lr IC'('5716 and l('('17101
LlST O F T t l E APPENDICES
Al)nendir No. I'agc No.
I . Sepregalion parrcrn 01'1:3 gcncrarion ol'thc crotr I('(' 5716 \ I('(' 17 101 75- 7 1
2 . Ind i~ idua l plant data of 1'2 gcncration o l thccro\ \ I('C .i710 \ I('C 17101 7 5 - 7 9
3, Individual plant data o f 1 3 generation cil'lhc crorr I('(' 5710 \ I('(' 17101 y[ , -g~.
4. Ind i~ idua l plant data ol'dill'crcnt gcnotlpc? tbr \$hitc t lo~ver colour Q-fY
5 \tlpnfhpr repon clilri!.:! tt!r: crop gro~vth period 86
LIST OF TABLES
--
Table No. Tit le ~ -
1 .Parental characters o f ICCV2, RS-I I. .I- 39- 1 and 'I'- I -A
2. Segregation ofF2 generation for stet11 pigmentation and
Ilo\\cr colour for the cross IC'C5716 s ICC 17101
3 Segregation o f Flgeneration for stem piglncntation and
I loner colour ti)r the cross I('(Y7 16 s I('C17 1 0 I
4. Joint segregation ol'stcm piglncntation and Ilowcr colour
li)r the cro1('('5710 s IC'C'I 7101
5. Crossing pattern ol'parents Ibr deterniination oSdiSSerent
genotypes tbr white tlo\\cr colour
6 . 'Table showing possibility o f crosscs among whitc I l o w c r ~
liotn thc crosses ol' ICCV 2 s 1'-I -a and US I I s '1' 39-1
7 Crossing pattern and segregation ol'wliitc Ilowcr types
from both c r o w s for getting triple recessive genotype Ibr
white Ilowcr colour
8. I'henotypic correlation coellicicnt 0 f 1 : ~ and I:, generations
of lCC5716 s IC('17101
- -- --
I'age No. - .- -- - .- . -
.i 1
i,
0. Phenotypic correlation coefficient among different \rhite
Ilowcr genotypes
1 con rest^ rr.rcrr2 rhr L o r d J l m ~ g l i t y f o r hrs 6rrrrrrdL,n bL,!!riri,$ r l fircli
or iompantedtnc rn al lendeavours
,D~itrorr 1s not crrnritjli 111 t r p n 7 r s my dcrp S P I I ~ C r I / l j r c r t r r r r~L3 ,trri/ / rirr11
t o place orr record my heariJd/ regard5 10 rry es/ccrrrr~d rliarnnarr if ,tdr,rsnry 1-i~rrrrnr/tr,e
iDr .yagdrsl i ' y u m a r , ,Pnrrcrpoi \ i renrrst, ( hrr.4,pca 6ncdr1itj /~rt~~nriitrorro/' ( nrpr
x~ , t r , i i r rh I r rs t r t r i t r~ j ~ r th1. Scmr A n d T~CJIII, I I ( %/.\.A?), ~ l + ~ t d r r c / ~ ~ ~ n t , .A ,I' . f i ~ r f i r 1
r n ~ p r n r r g orrd i I / ~ c / r o ~ r o t r ,lutdorire ~ ~ ~ r r s t i z r r t i ~ r r r i r i ~ n ~ ~ ~ ~ r n ~ ~ r ~ ~ <lrrd c ~ r r \ t n i i t r r ~ . rn t r r r r rn
r r r phtrrrrrng ~rrd11rr,r t , rr t t r t r~~rr o ( t f i ~ , rr,~r,urccli r r or(
I om p n r f i u ~ ~ d l y rrrdchtcd t o (Dr. ' 7 : f l a g e s h w a r qpo, '~)nl /r .ssor , r r r d
/ ( rod , ~ D e p i i n m e ~ r l 19 (,errcr~is arrd P h r l l ~Hrcr'dtrrlj. (r l l lc i jc i l / . A g n c u ~ t u n . 'Y<t /~~r rdrd
rrogar os ro r l ia~nni i rr (I/ my art [ r r q rnmmrt t rc j r r f i r 1 rnunljlcrlrr/ ' r r i j r rre~icrrrc i rnd
rncttcr ih~ur rr9~rsorrrrr~] 111 rej~rrp thrs d ~ r . ~ r ~ r i n t t o ~ r morr e~p i rc r r ry t n rr~c[orr rr,rth u s /
rtandards lrrefjubh tr my j]~1trtrr&2 drrd srrrcL,rd tharr(r t o film / o r f i r i / rorr\rerrdcr~t
s~r ,q ,qc~t~orn 110 cm6elhrh the ,A/ rrdy
I ' f i r tny pr~$ise t h a r r k ~ 111 ( i ) r . \ u b h a s h r t i a n d r a , ccrrtor .ccrctrrrlt,
\ la t rs l l rs , I r r t c r r r a ~ r o ~ ~ a ~ ( ' ~ r p r 'Yertcrrrli l r l s l l l u l c j i l r the $ern1 / Ind?nrpr r 7 ('I('yl,SJ'1],
~ I ' I I ~ N I I C ~ C N , j l 1P as rrrernfier of rny n d ~ , r s o ~ commrttcc j r r r fils g o o d r o r i n ~ c l nrrrt help
h i n r r g my course ~ftrrrr2strga/rorr a116prepara11~11, rd/I;err.r
I t 1s my pnde arrd hrlrrour 10 erpres, my prr!firrind i ~ v t t c i f l j r (zt1rude arrd srrtrere
r ~ y a r r i r 10 / f i t . 1n:m6t,r r l / odrtrnry coinmi/ tcc (Df L !M Fpo, ~~'nf i .ssor, ,l)i.par/rnent o j
(Plarrt ~phyctoiogy, ( o l L y e rf A g n r u i t u r e , ~~ai(alerrdrarragar J n r fitr arncr~aljlc nrrd
s/uperrdous gurdarrrr arrd c r ~ r n u r a ~ e m c r l / arrd ua luahh ~ u g g r ~ t r r ~ r i s 111 quaL la t tvc
tmprovemenl Lo the t e a
I w o u l d bke t o thank a r ( L L G o w d a /{cad; L .\ O' I( KI.Yj'1.
Patanchcnr fo r pennrtt lng rnc a n d p r o v r d ~ n g f r c i d ; l a b u r a / o r y j a r ~ l ~ t r e r arrd roopera / r~r r
dur ing t i ic course r fmy tnvcsttgatton
I am t f ianv~/ l r { t i? Mr. (H.'L! ~ Q O , .Sir~,rrt!/ic (
INHERITAhCF O F S1 EM I'I(;MEN r,\I'10\ ,\\I) FLOWER COLOIIR IN CHICKPEA (Cicrr crrirtir~rrrt~ I
hIASTEK O F SCIENCE IN t(;RIC'l'LI'IIRF:
I o \lutl! rhc ~nlicr~r,~ncc 111 \ten1 plgrncn1,lllon ,inti Iloucr ~iiIo11r I( C 57 1 h '111d
I C C 17101 \rere ujcd 37 purcnlv 111 \v l i~ i l i I( ( 5710 %,I\ \ r ~ l h Iilgh p~gnic~i lct l +1cni ~ n t l
p ~ n L llot\er ,~nd IC C 17 101 %,I\ u ~ l h Io\v p~gliicntctl ~ l c m ,~nti \vli~tc Iloucrcd Inlicr~l,lncc
+ lu t l~c j ucrc 11i,ldc 10 stud! slcm plgmclitatlnn. tlo\\cr iolollr and rl\+O1ldllO1l jludlc+ lor
tl~llcrcnt quantltatlve characters
hlonogen~c ~ n l i c r ~ l ~ l n ~ c \\a+ conlirmcd for llic I\ro rnorpholog~c~~l ~ l i a rd~ I c r \ I e
10ir plg111c.ntcd i \ li1gl1 p1g111~11tcd ,111d p~nL Iloi\cr LOIOLI~ 1'\ u l i ~ t c Ilo\vcr LOIOLI~ I OLI
\Icm pglnentatlon dorn~nalcd 111g11 \leni p~g~ l i cn l d t~ (~n charilctcr and l i l \ \vcll 10 lllc
cvpccled rd lo i I I'lnh llo\vcr colour domlnatcd the u l i ~ t c l loucr colour and l i t \ uc l l lo
the cvpccred r,irlo 3 I .lo1111 +cgrcgallon 01 ~hcsc r\vo character\ +lioucd ~ndcpclidcnt
a\\ortment and fits uel l to the eupected r,illo 0 3 i I ~ndlcal~l ig lli,~t rhc\c ~ u o c l i ~~ r~~c rc r \
arc iolilrollsd h! I\\(, d~lferenl gene\ I hc results (11 I gcnerdnx \rere colilirmcd 111 tlic
\rudy of I 7 generatlnn
111 hot11 I:? and l :~ gcticr,1tio1is !icld per pI;~rit \\;IS corrcl;~tc~l \\it11 pl;ttit l i c ~ g l i ~ ;111tl
n i t h nutliher o f pods per pl;uit. til~tiihcr (11' seeds per pl;ltit ;~tid t i l~~i ihcr 01' hcco~id,ir!
hranclies per plntit. 111 [:: gc~icri l l io~i ~ i l~ t i ihc r o f pri~iiar! hr;~~iclicc per pl;ltit ;~tid it1 I:, geticrotioti tln!.s to tiiaturi& ;itid liltlidred seed \+ciglit \+ere 1101 ;~sst~ci;~tcd \\ttI i ;tti! 01 tlic
clinrnctcrs sr11dic.d. Kotiiher o f sccds per plant in boll1 I:: inid I : gc~icr;~tions pt~\ i t i \c l !
corrcl:ltcd \\it11 plant Iie~gllt. ti~ttiihcr (11' scc~tiiI;~r! br;~~icIics per pl;t~it, pli~tit \ \~d t l i . illid
icld per planr.
111 c lcrcr~i i i~ i i~ ig dil't'crc~il gc~iot!pcs for \\liitc l lo \~crc i l l![w, I('('\: 2 , I t s I I (\\ l i i tc
Ilo\\cred) ;lti(i I ;')-I. I - I - , I ( h l o ~ Ilo\rcrcdl \\ere ~tscd ;IS parents. ('rosb hct\vcc~i the \\Iitrc ,111d hluc I lo\ \crc~l p;trctit\ (I('('\.' 3 >. I - ] - , I ~ t l i i l ItS I I Y I 30-1 llicir 1 ' 1 \+its pink i lo\\cr 111 hc~tli c;l.;es atill tlicir I,? g c ~ i c r ; ~ t i ~ ~ n hcgl-cg;ttctl as pitih. hluc :lntl icliitc. l i ls \\ell
tu [lie r,tlio ol'O.?.J it i i l icnli~ig \upplc~iicnl,tr! gcnc ;tclto~i I lie gctlot)pc\ Iilr tlic p;lrclilh
\\ere ~ l c ~ c r t i i i ~ i c d as I('('\: 2 ( '( '/ilil'P, 1'-I -:I ( '( '11111~1), 1;) lpi~ih ('( 'I1/il1/) 1': liitik ( f : h l I / i d i i t r - I i l i l - t i - 1 1 1 1 (ictiot>pc\ 01' I t s I I
L,L 11171'f. I 30-1 ('('1111/)/1, pitih ('~11111'~~. I:! lpi~ih ('-11-1'- 1,: l i l ~ tc ('-11-/1/i I ?i+Iiitc
1.c /!-IJ- ;111d C,C 11-/l/l
I3! ititercro\sitig tlic ahtnc t i i c~ i t i~ t i cd icliitc I10\+crcd g c ~ i t ~ t ! ~ ~ e < [ ( I ( 'CV 1- s I - I - . \ 1,; \+I i i~c llo\vcr) (l<S I I i I' 30-1 1,': \\Iiitc l lo\\cr)! i l l I,', 1pitiL ~ ~ t i c l hI11c Ilo\\cr
\\ere rcsultcd. i l l 1.2 pink Ilo\rer 5egreg;lted itito pink. Ibluc ;ind ~ r l i i l c ( ' ) : I4
\upplc~iie~ilar! gc~ic ac~itrti). hluc ilo\+cr \cgrcg;~tcd intc~ hluc atiil \+liitc (0:7
co~iiplc~iic~itar! gene acttoll). In hotli case\ \\Iiltc Ilo\+ers ncrc rcsultcd atltl their gcllot! pch ncrc i i \cd a\ \+ere ( '-hh 1'- 1 . ~ 11-1'- L.CW ,+ ( '-/I/I/I/) ~ .~.hhl ' - i .~.hh/~p 111
this s~ud! ;I tripplt. reces\i\c gc~ic~t>pc lor \+li111, ,,(.i\cr 1.c.. cchhpp \*a\ dclcrmi~icd Iiir
rlic fir51 l i~ t i c .
,Inlong dil'l'erctit \+hitc flo\+cr gcnntlpc.; l i c l t l per platit. slio\\cd ;I ti011 sig~iiIic;~tit
as\ocialtoti \+it11 all tlie traitr ~ttidcr \luil) csccpt \+itli. ~ iu~ i ihc r o f pod\ per plant. Nu~i ihcr
( ~ i ' pod\ per plat11 pc~\iri\cl! correlated sigtiiiicantl!. \+it11 plant Iiciglit. ~iumhcr 01'
~ce011~13r! h r ~ i n c l ~ c ~ . t i l~~ i ihc r ol '~ccd\ per plant. plant uit lt l i ;lnil ! teld per plant.
DECLAKA'TION
I. KlRAN KUhlAH V.S.S Ilcrehy dccl;~rc h a [ [lie rlichij
cnrirlcd "INFIERITANCE O F S'TRM PI(;bIENTATION ANI) FI,OWER
('0LOL;R IN CIIICKI'EA (Ciccr aritfinirnr I,.)" suh~iiiltcd Lo ~Ic l ia ry ;~ N L i .
K;lrlg;~ ,Agricultural I l~ l ivcrhi ty for thc degree ol' Mi1I;'I'I'l~ 01: S('II:N('II I N
! \ ( i I~ lC ' l~ I , ' l ' I IR1: i ? a resulr of original rcwarch \\ark dolie hy Iiie. I a l ~ o declare
11iat tlie Ilic\is or part rliercol' 113\ nor heen pnhlislicd carlicr c l~ewl icrc in ;my
Inanner.
LIST O F ABBREVIATIONS
100-seed weight I ISW
('entirneter ~ 1 1 1
C'hi-sqi~are 2
I h y s to 50% tlo\vcring 1)1~50%
Da>.s to tirst l lo\vcring 1)I~l.'
I)o),s to first pod tbr~nntion l)l,'l'
Day\ to t i i a t~~r i t ) I)M
I:Io\vcr COIC)II~ I:('
Grams 1:
I lcctnrc ha
Mi l l i on 'l'onncs M'I'
Number ofpodc per plant Nl'f'
Nurnbcr ol 'pr imary branches per plant I'b
Number of secondary branches per plant Sb
N u ~ i i b c r ot'seeds per plant NSI'
I'lant hciglit 141
Piant width Wd
['robability P
Stcrn pigmentation S I'
Yield per plant Y ld__l'
l ood Ic.gume\ pl,n .I m.ilor role In o g r ~ ~ u l t u r c h c ~ o u ~ c the) arc an ~rnpl~r tant
\ourcc ot food ,lnd ~ d p ~ b l e ol l islny n t m o a p h c r ~ ~ nltrogcn through thclr ,lssocl,itlon \ \ ~ t h
Ritizoh~rr h . ~ c t c r ~ u m ~onscquent ly they can be yro\\n In \ o ~ l \ ol low tcrt~llty. and they ,ire
rclerred to ,IS mlnl lc r t~ l l rcr tactorlcs Racy ut~l l /c 5011 rnolsturc clliclcntl) h c c ~ u \ c ol
their long root\ and have c a p a ~ l t y 01 b~cldlng \onic th~ng c \ c n under rn,lrg~n,~l , ~ n d 1110\t
~icglccted ~ o n d ~ t ~ o n \ ,rnd w ~ t h Io\r Input5 Ihu\ they cnh,lncc \ O I I I c r r ~ l ~ t ) 'ind c,ln he
g r o i \ ~ i 011 u ~ ~ i \ c r v c d rcs~dual so11 ~nio~\ turc I Ihc) aI\o 111ipro\c \o11 p l l ) \ ~ c ~ i I \tructure
C h ~ ~ h p ~ d (( ~ L L P rlrielinllm I ) I \ c,~IIcd I3cligdl grlllii or grd~i i 111 Ilidld and
garbanlo hedn In m u ~ h ol the de\eloped world It IS the \\orld'\ thlrd mo\t Important
food lcgumc crop In ternis o l ~ o n a u m p t ~ o n and b) tar the most Important p u l x 111 \out11
Asla I he \ e r s a t ~ l ~ t y ol c h ~ c k p e a In culslnc 15 Icgendar) (Vandcr m,le\cn 1072)
( h1ikpc.a Ir currently grown on about 12 01 rn ha worldwldc In ~ h l ~ h Itidla lid\ 'I \hdrc
of 69 Sola w ~ t h 8 4 m ha It ranks first w ~ t h a share of 72 8% In the \\orld production 1 hc
p r ~ d u ~ t ~ ~ ~ t y of chlchpea IS lo^ as comparcd t o ccreals. r ~ n c e chlckpcas are g ~ n c r a l l )
gro \ \~ i in poor-soils and droughl-prone, r~iv i ron~nc~l ts. I'lie \\orld's 111cnri productivit! i h
768 hg:11;1. India's p roduc~ i t i~y is 708 kg:'lla (I'AO. 2000).
Sclcc~ioll pla>s a vitnl role ill \ucces\ trl' a breeding prograniliie. \\liicll dcpcnd~
up011 rllr nalurc and ~il;lg~iitudc CIS i~ssocia~io~i bctwccn clinrac~crs. So. lo l i l id out 1l1c
;I\SOCI:IIIOII. cl~rrclatioli I\ ;I t001. \\liicIi cnn he used lo idcntil). llic uselill cli;~rnctcrs
i~ i f luc~~c i r ig hiclil nil ~milcsirahlc ,~ssoci;~!cs hcticcs~l tllc conlpollcnl cllaraclcrs. I'liis
crilpliasi/cs tlic importance ofccirrcl;~litin at~ldics.
I'lic (1st o f ~iiarkers i l l a crop cultiv;lr gibes ;III addcd od~a~i lagc i l l
cllnractcri~i~lg, sclcclion ;~nii ill mi~int;~ining tlic gc~ictic purit!. Scbcrirl ~iiorplic~logical
~il;~rkcrs l i ~ r shape, colour, sire. ;lnil pigmcntatiori arc used in dillkrcnt cropc. 1:loircr
colour nnci stem pignlcntntion arc useful ~llcirpliologic:rI ~iinrkcrs in cliickpc:~. (;c~icticb 01'
\aricrus Ilo\\cr colonrs ofcllicl\pca i \ 11ot ircll understood a5 ;~lso l i ~ r 1ii;111y otlicr tr;lits.
I lie av,~ilahlc inli>r~ii;~tio~i on niarl\er\ 2nd traits 2nd tllcir l i~ik;~gc is rather liliiitcd. I Icncc
tlic prcscnl sludy is conducted wit11 [lie following oh,icctivcs.
I , f o investigate tlic irilieritancc o f stcm pig~ncntntion a id Ilo\vcr colour.
2 , I o detrrniinc different gcnotypcs Ibr \+hitc Ilowcr colour, and
3 . l o investigate association among imporlant traits.
Revie\\ o f literature pertainilig to inheritance o f stcni and Ilti\*er pigtilcntntion.
I lo\\er colour and correlat io~i al l iol ig different traits i n cliickpea (('ic.o. tr~ielirrrrrir I,.) is
~'reselited briefl! ullder the I'ollo\\ing Ilcadings:
2 1 l ~ ~ l i c r i t : ~ ~ i c c o f s t c ~ i ~ :!II~ l l c ~ n c r p i g ~ i i c ~ ~ t a l i o ~ ~
2 2 l t ~ l i c r i t a~ i cc o i llo!\cr ccilrit~r,
2.3 Associat io~is al l iol ig i ~ l l p o r t a ~ ~ t traits (('crrrelatioli coeI'licicnl<)
' I l i c s t c ~ i i atid I lo i rc r pignientatiotis ill cllickpca are usel'lll ~norp i iu log icn l
11i;irkers
I3llalikar arid I'atil (1062) l i iund Illat foliage and I lowcr colour arc governed b y a
single Iclclor i ind a l ~ n rcp(~rtc(I lliat tlic sccd coat colour *as crintrolled by t*o
co~i ip l in ic l l la ry Ihctors. I lic) furtlier po i~ i t cd c~ut tliat tlic factor rcspo~lsihlc li,r foliage
colour was dif ferent and indepeudent froln the factors re~pons ih le Tor flo\cer and seed
coal co lo l~rs .
Al ier slid I'atil (1984) studied the colours o f steln axil, pedicel. cnrolla and veins
011 [lie ventral surface o r t l i e stalidard petal. I l e reported Illat all tllesc trait? ;Ire ctiritrolled
h! a single plciotrop~c gene: I.I.(.o This gc~ ic is i ~ i d c p c ~ i d c ~ ~ t llic ~ ~ I I C S l I r , \~.o a110 )'\(,o
that control seed testa colour.
N;I>u and Rhat (1984) fiiund lhal colours oI 's!c~n and corolla ill coffon \\ere
ct>~itrollcd h) t\ro genes and one gene arid rli;lt 1Iie red p ~ g n i c ~ i t ; l ~ ~ o ~ i \\;IS donlinan! 111
stclii, boll and hrnct and i n c ~ ~ i ~ p l c l c l ! do~l i i~ ia i i ! i n tlic cor~il la.
(iliatgc and Kolllc ilOX5) reporlcd tIi,~t pmh i,\ \ r l~ i t s colourc o l ' s l t . ~ i ~ , pedicel.
scp:ll, pc~n l ;lnd pclnl vein irere Ibund to he gobcrned hy ge~ics ;Ir t u o loci. Il'corr and
H'coh \ \ ~ t l ~ p ic~otrop~c ;~ction and tlicsc colours s h o ~ c d all 1'2 ccgrcgarion ratio 01' '):7.
I~sso~ i ih ;~ rri (1087) ~ l i o sludicd I,'? co~iiplctc dlallcl cross ol' pconul suggcslcd
lliat purple and greeri p ignie i~~:~l io i i were ccintrcillcd by at lcasl onc and two duplic;~lc
gcncs illid epihti~tic aridt(~r additi\jc cl'l'ecls cxihl l i ~ r growth arid inter nuclear Iilctors
~nllucncc relationship hctirccn traits.
'l'lie sterii pigmentation depends upon prcscncc or ahsclice ol 'arit l iocyo~iin
pigmcnt. uhich ic l ig l i l dcpcndcnt Llalhur (1080) reporlcd a cliickpca line, which
debeloped purple pigmentation in thc irholc plant, stcni, hranclie. Icaves and I l o \ r e r ~ arid
concluded that this purple pigmentatii~n is utterly light dcpcndcnt. I l e reported lhat tlie\c
plants do not produce pigmentation if not directly cxposed tu sunlrght.
Rahir and Sen (1991) fuund that nl i t l ioc)nn~n plgnicntntliin o1'1lic s tc~i i ir
controlled by two penes. one oS\\liich sliowcd partial dc~riiinaricc slid \\as cpistotrc 111 the
cillicr gene in thc recessive honiozygous state.
Karkan~ia\ar el ill. ( IOOI) proposed drgc~ilc inlicrltancc ofs tc t i i pigmclitat~on with
co~nplc l i ie~ i~:~r ! ycnc ,letion :rnd a 17.4 %, crosso\cr \ u s see11 hct\\.ccl~ onc
cornplc~iicntar) gclic l i i r \ ~ c t i i pignicnt;~tion .111d l l ic ge~ie li)r stipirlc colour.
\let/ el 111. (I')')?) studied ~nhcri~:lncc ol 'pt lrplc sccdling colour (I'SC) and its
rclatioll 10 gclietic ~01itro1 11s I l u \ ~ c r CIIIOLI~ It LI,I\ l i l u~ id t lu t I'SC' is prohnhly colitrollcd
h) :I singlc gclic. I'licy Surtlicr (ib\ervcd that this trait is diinilndnt iivcr grccn cccdling
culour I-lic ~hl i i tc t l i i \rcr i r l l i l h l t~d tile cxpres\ i~ l i ol'I1S(' and \has tlicrcliirc l l iougli l to he
cpislallc.
. I l i ~ i i ed slid 'I ntiki (1002) reported l l ic y ~ r t l a l doliiilialic(~ 01' purple \ t e~ i i o\er
grccn \lem in chi l l i ;rnd cih.;er\ed contiliu(iu\ \:lrlatlcin in I ? :rnd tlic hack cross
generottons indicated polbgenic control ol'anthocyanin pigmentation.
Sandhu el ill. (1Y93) obtained I 3 green tollage and 3 purplc l i ) l~agc In 1.2 alicr
cro.;sing purplc pigmented plant as male with ~ iorn ia l grcen l i~ l iagc. I hc) ruggcstcd
digcnlc inhcri~ancc o f colour \hit11 dom~nant and rcccbsi\c cplstallh In 1.1 nornial grccri
Ibliage ia dominant o\er purplc foliage thu.; thc plants n i t h 1'-I-/l!ljl- / /'-!I gcnnt)pcs
have grcen foliage and those ~ i t h the P-ii genotype have purplc liiliayc.
Singh e / ill (1993) studied FI. I:? :111d I:3 gc~icrdlic~tis hy ~1si11g t\%o pigc1111pc;t
tarielies and l ive ICP lines. I lie t u n IC'P lines lCPl 1104 and ICP 8802 Ii;id purplc atem\
and rcniaining ;trc of green. 1111 I:?s scgrcg;~tcd ill rlic r,trio .? purple : I green.
I:3 produced 1 purplc : ? segregating . I grccli. \~~ggcst ing tI1:tt stcni pigniclitntion i.;
controlled h) a s~nglc gcnc \\i l l1 purplc as doniin.itit.
(;Ii;ttgc (1'1'1.3) studled tlic crosses 111 cliiclpca rcvcalcd tIi;11 cotylcdr~n colour
\\a\ cr i~ i~rr i l lcd h! a ainglc gene (li.i~/). u l i i tc Iiiliugc cri lol~r \%as duc to )io/ and li1,ylo.
(;li.ngc (1994a) indicated tliat purplc stclii crilour IS doli i i~iant obcr grccli stclii
colour aliil ptrih curoll;~ domin,~nt over ivhitc. ' lhc gcrlcs govcrllitig lllc ilillcri~nncc ill'
,tern colour and corolla culour are designated as 1',/ ; t~id llL.O itlid MCil and Pcl1
rcspcctivel) with n,,, being common ro boll1 trait\.
Joslii P I (11. (1994) reported the existence 01' ;I hasic plciotropic gcnc (1'1)
rcsponsiblc Ibr thc cxprcssion o f pigmentiltion of itxil, calityx. corolla, pod tip and wed
rogciher u i ~ h locali/ing genes conditioning colouralion 011 \pccilic plant pans In coupcjt.
Alimad (1947) indicillcd hat the stem colour i j dctcrtnincd hy Ilic intcractlon 111'
~hree pairs genes namely P-P 1-1 and R-r. P-p (green pigrncnlatir~n). 1-1 (ciilnllr
intensity) and R-r (plgrnent reduction) i n kenat:
I'undir and Kcddy (1097) reported a natural niutant \\Iiicli co~iihine.; pl~rpl lsl i stclll
( low p~gmented) with white I loaer. 'This trait conihinntion \+as not pre\iously L~ iown ill
( ' i cc r and designated this niutant as ICC 17101
coniplinicntary in tenct io~i o l ' t ao genes ( O : ? ) fhr light depe~ldent purple ( l . l) iJ) and noli-
1.111' pigmcntcd pla~its h) using IC'C' 12 ; ~nd I( C 10.301 parents.
Ve11ugop;ll ;itid (ioud (1098) rcpoflcd Iron1 [lie segregntlnn n11;il)sis 1Ii;it trll
pignlcnlcd character\ wcrc d o ~ i i l n a ~ ~ t o\cr 11011-p~gn~cnlctl cl1nr;iclerc in c o ~ p c ; ~ . I lie 1'2
pl icno~)pic ratios oi' 11:s ( two tlircslicilil diiniin;int gcnc.;i. O : ? ( two co111plimcnt:lr)
gcncs). 3: I (nionogcnic doniinant). l l:5 (two tlircsliold donii~lnnt gc~ies) 15: I ( t a o
duplicate gcncs) and 54: IO (an) two 01' tlic co~iipl i~i icntnr) gcncs) wcrc observed liu
plgniclitatloli on seco~itliir! and ~crtiar! hr;lnclics. pul\inus, at;lll\ tip. pctluliclc surlicc.
peduncle l i p and stipulcs rcspcct~vclj
I'cfcra (19'18) conlirnicd monogenic ililicritancc li1r tlircc morphological
charnctcrs pink v.v white I lo~.ers, pigmented Y \ non pigriie~ited stem c o l o u r ~ :~nd w g l c
poddcd ~v double poddcd characters. 'The flower colour 01' gcnotkpc IC( 'V 2 \+as
dctcrmined as I'l'hh('(' and JC; 62 as PPI1R('( '. I his gene \*as h u n d to ha\e plciotropic
cfl'ect as it conlrolled the vlem colour as \vcll.
Sahaghpour (2000) studied the interrelationships hetween pairs ofcIi,tracter.i sucli
as: tlo\\er colour. stem colour. seed coat colour. I l e ohserved that the pcnc h contn~l l ing
the Ilower colour had a plciotn~pic cl'kct nl i steln colour supgcsllnp Inonopcnlc
~nhcrltance of Ilower colour patterns in chickpea.
Single gene inheritunce model was proposed hy Met7 o (11. (1007). (iliatgc
(1903). Sinph el (11 (1093) and ' l 'ckr i l (1008). dlgcnic inlierit;~~icc 111~1dcl \*;is proposed by
(ihntgc and Kolhe (1085). Mathur (IOXO). Kilhir iind Sen (1901). Sandhi1 c/ (11. (Ic)Oi).
N;~hu ;lnd I3liat ( 10841 and trigenlc model \+;I> proposed hq A l i~ i i cd ( Ic)')7) ;lnd polygelilc
~nheritancc \+as proposed hy i\hmcd und '1';lnkl (1002). I rum llic ~ h o \ c m c l i ~ ~ ~ ~ n c d
\rudiec the stciii and flo\\cr pipnicn~atinn in chickpea arc reported to he n~otiopc~iic.
dlgenic or tr~gcnic r1ict.i at least three genes go\crn rllc \ten1 pigmeli~:iticin ;tnd I lowcr
c o l ~ l ~ i r ill chickpc;~
2.2 INIIEKITASCE OF FL.OWI<K COl.Olll<
In chickpea. ~herc are tlircc distinct t loucr colours naniely pink, hluc and uti i tc.
I \*o uhl tc llowcred varieties 1' 4623 and IKS I I uhcn crossed produced pink Iloi+cred
hybrid indlca~ing 11ic presence o f diflcrcnt gene9 ct in~ml l ing lhcir Ilo\*cr colour (Kumar.
1997). The sludy o l f loucr colour inheritance in cliickpca is import an^ in crosscs
involbing such parcnts, 11 re\lew of literature fbr I louer colour lnherilancc In chlclipca is
presented hereunder.
Khan and Akhvar (1934) studied gcnctlcs o f t l ~ jwer colour In 1 .1 and 1'2
generalions o f xvera l crosses involving blue, plnk and \ \ h~ tc fluwcred chickpea typci.
l'he! reported [hat [he hlue ccilour \\.as due to a single thclor 8: a Ihckir I' g;nc pink
colour in the presence o f B hul \+as hy ithclf \.ritlliiu[ coliiur ct'l;'~~. !I green colollr 01'[Ile
standard petal u n s ohmined in the ahsencc oi' ;I hcki r H' I'hc! oh[;lincd a r;l[ici 01'0 pink:
3 hluc: 4 white colours in crosses involiing \+llite and blue t)pcs, i ~ n d 3 3: 1 1,1110 o f p i ~ l k
a i t l l hotll blue and \\liite. pink being dollli~lnnt lo either. I .n~cr I':rl (103.1) ohtn~ncd
sinlilar results based 011 a series of crosses ;rnd conlir~llcd 111at tlic I10\+cr C O I O L I ~ was
governed h! rile i~l[ernctici~l cift\ro gcllc pair;
.I!y;lr ;111d IIalasubra1l~a1lii111 (lO3hi to~111d II1;11 llle i ~ l l l e r ~ t : ~ ~ ~ e e 01' l lu\\cr C O I O I I ~ S .
pink. blue and \rliitc \\;IS go\cr~lcd h) three L~clors. 1\40 conlplc~ncnrar) ihctcirs ( ' a n d M
\+llich together produced hlue and ;I t;~ctor I' \\llicll co~l icr tcd blue 11110 p ~ ~ i l \ . 111 the
ahscnce o f ( ' o r 11, the I loacrs a c r e a h ~ t c .
l'iniplikar (1943). Khan c f trl. (1050). f'ntil (1004) and i!tll\+al and Ilrnr (1067)
studicd crosscs bctaccn \\hitc and pink tlo~vcrcd strams. and ohscrvcd [hilt the pink
colol~r was dominant to a l l i te Ilo\+cr. 'I he) obtained pink and wllite Ilo\.rcr segrcgatioll in
the ratio or 3 : I
llliapkar and Patil (1063). Patil (1967) and Na)eclli 1.1 (11 (1077) s h o a c d from tllc
crosscs between blue and pink flowered mutants that llowcr colnur \\af rnonogcniwll)
controlled, with pink being doniinant to hlue.
D'Crur and rendulknr (1070) shoacd liom I:I and IF2 gcncr.i!lons of the cross
Douhle pod s H'h~ tc flo\\er W h ~ l e grain that tlircc gcncs I'cii,,. I'ciih, and I'r.i),,,
go\crncd corolla colour and a.eri: pleiotropic in controlling stcm and coroll,~ colour. I'licy
detected llnhagc hct\cccn corolla colour. nuniher ( i f fl~i\vcrs per axil. Ics l i~ colour and cecd
shape.
Khosli-Kliui ;~nd Nlknqad (1'1711 and MI;III (1071) studied I:], I:? :lnd I:3
gcncr;~tions o f reciprocal crossca hctwi.cn \ \ I i l ~c ;111d purple Ilo\bcrc illid ohser\ed
non no genic inhcrltance ( i~ r ll(i\{er colour. purple hung ilom~nnnr o\cr v.lii~c a ~ t h no
malcrnal ctfcct\.
I'hadn~s (1076) conducted ~nlicritancc \tudiea o n I:? and 1'3 gc~icrations o f
cniaacs hct\\ecn lines ha\ ing \vliilc or pink Ilowcra i n f 'icer oric1117iim. I lc noted that /1
\+a,, .I dominant gcnc for plnk Ilo\rcrs. 11 and ( ' c;icIi slngly rcsullcd In \\hite l l ~ i a c r s hul
ucrc complcnicnt:~r). resulting in pink ilowers \{hen both were prejent: one o l rhc
complcmcntar) gcnea lnhibitcd , I .
Reddy and Chopde (1977) studied t\co crosses in ( ' i ~ , c r irr iel ini~n?. In a croaa
hct~vccn \ iolet f l o ~ ~ c r c d Chikodi V.V. and p ~ n k ilov.crcd ( 'hry\unlhefi~i io type. two
complcmcntar) gcncs, dcslgnatcd I'co,, and Palh conditioned f louer colour, pink being
dominant. In another croaa hctucen \ iolct ilo\\crcd Clilkodi V V and \\lilte I l(~\\crcd lypc
Kh. 908-21. thcy obser\ed that a alngle dominant gene Lvro governed Ilov.er coluur.
violet being dominant.
Kaii 1.1 111 (1980) studled fhc ~nhcritancc ol' l lgl i l blue corolla in 1.). 1'2 :uid
hackcross generations 01' scvcn crosses tnvol\ lng hluc and p ~ n k Ilo\+ered t lpcs. Ihcy
ohrnincd pink IY l s and I:? segregation rafiil o f 0 pink -3 l ~gh f pink : 3 hluc : I light hluc
I'lie) sIio\\cd flint I~ghl-blue corol l :~ ~nvolved Intcracflon ol ' tuo rcccssivc alleles.
Kumar er '11. (IOX?) studled I.'? scgregatlon r;ltlo 01' Ilic cross. I-I-,\ x Annlgerr
arid ~nd~cn lcd flint Htlwer colour \r,lr n ionc~pc~i~c:~ l l ) irhcritcd. \ v~ l l i pink hcing d ~ m i n ; ~ n f
I l luc Il<i\\crcd pI,lm\ Iiad Ihiglicr secd profcln confcnf ;~nd \ni;llIcr sccds flian pink
Il~iu.crcd plants indicating l~nL;~pe hef\rcen l11c genes goberning Ilic Ihrcc ch;~r;~ctcrs
1.1nknge was not t~g l i f , liencc fhcy conclutlcd t l i ; ~ ~ scgrcgaflng pl.lrna cumhining a l i ~ g l i
seed protein percentage u l t h I;~rge seed\ niiglit he rcco\cr;~hlc from I:lrgc pi)pul:itions
l'audr atid I'atil (1082 and 1083) liiund In the cross. ( ' l i ikod~ V.V x 1)-70-10 fliaf
m g l c gene I'cri controlled corolla colour, with plnk dominant to light \ iolcf. I tic I:~cfors
Iur c~irt i l la colour. seed coil1 colour and sccd surtjcc Sorniccd onc Ilnkagc group
K ~ d a m b ~ er u!. ( 1988) atud~cd parents. 1.1, I:?. I3C'l and RC? gcncrallrinh o f a
croaa hel\+een \+bite and purple Ilo\+er type\ f lo \ rer coli)ur !eprcg;~tion ~n 1.2 and
I<(') indicated that purple was monogenically dornin;lncc ovcr \ rh~ tc ciilour
Singh rt ol. (1988) worked ouf the association among I.usarium \v11t revstance.
flower colour and number of Ilo\+ers per fruiting node In c r o w s made betuecn
genotypes differing for the above characters in cliickpca. They observed that h ~ d
pink flo\vcrs and rhe ratio o f pink to white Ilo\vcrcd plants i n F2 \\as consistent u i t l i
segregation o f a single locus. h i l l1 pink doniillant over \vliitc. 'I'hc) observed tIi;~t tl(i\rer
cnlour \bas inherited indcpcndcntl) o f t louer nunihcr nnd \ r i l l reaction.
Da\.is (199 1) in~cstigated [lie l i~ihage rclarionsliip o f gcncs Ihr Icaf ~i iorpl iolog).
tlo\rcr colour and root noduliltion in crusts betireen purple and white Ilo\rcrcd lines.
and ,Imong u l i i l c Ilo\rcred lines. I l e dcm(1nstratcd that l l ic l \ ro \rliitc i lo\rcrcd lines
c ;~rr~cd ion-allclic. single rccc\.ilvc gcncr lor u l i ~ t c I lu\rcr colaur. provisi(11iall)
tlcsignatcd 11 1 end ,I? respectively. I It. shoired that tlic gcnes li>r l i l i l i ~ r ~ l ~ leaf tr:~it /il ;lnd
it'? \<crc IlnLcd.
Stephens and Nickull (l1)0?i Ibund a ~ i c u i loucr colour pink i n soyhe:ln. 1111
\el l ing tlie plrik Ilo\rer colour is controlled by a single rcccsbivc gcnc. ir l iun crosrcd u i t l i
all reported t loucr colour gcnes, tlie pink flower genc ( I V f ) is independent o l 'a known
i loucr colour genes and acts on nioditicd gene.
(iil and Cuhero (1993) studied the rclationsliip o f \ccd coat th~ckncss to \ced s ~ r u
and flower colour in the crosses betueen pink tlo\rered des~ and \rliitc Iloircrcd kahu l~
t l p e u n d obtained the cxpccted ratio o f 3 pink: I \rhitc u i t h pink dom~riant w e r u h ~ t c .
1.1nkage wa5 found hct\recn seed coat th~ckness and t louer colour, thc rccomhinant
fraction being 0.19.
Raini cr ill. (1994) reported that gro\\lh hahit tlotrcr colour, pod colour. pod
shape and seed colour are each controlled by t h o genes. Singh rr 111. (1094) studled 1:).
F2 and F ? of pigeonpca croshcs and suggcstud tliat red purple Ilo\rcr colour nos
govcmcd hg a singlc gcne uilh complete dotninancc o\.cr yclloa colnur:~lio~i.
Singh cr ill. (1944) itidicatcd thnt red pi~rplc I l o ~ c r colour \\as govcrncd h) ;I
single gcne \rith ~ncomplcte domin;lticc o\cr ycll~nr cnlol~r;~tio~i tIi;~t poll colour \KI$
go\ernt.d h) a single gcnc tvtlh coniplctc domlnancc.
S~ngli and Singh (1005) Ibund that tlic prupctiy ol'tlic irolct u aliitc ~.roaauh
liad violet tlo\vers. atid in the I:? the ratio of violet lo uliilc Ilo\rer\ ails 3: 1 wggcsting
tlint \lolet tlo\rcr colour u a \ goicrned h> hingls doniinont gene.
I l u i \ c d ~ ci 111. (1940) rcpor1t.d in groundtiut an unrti~hlc aliite Ilo\vcr colour lion1
(lie crobs hct\vcen t a o yclloa lloacrcd p;~rcnts and coticludcd tlial the prohahly sourcc
fhr thir inconsi5tcnt segregation for llouer colour is the presence of at1 utistahle genetic
element along uith the alleles producing trhitc floaer phenot)pc.
Venupopal and (ioud (1906) reported horn I: ] and 1.2 tlic calyx pignientotton
\vah controlled by three duplicate genes in coupea. Violct floacr colour bas dominant
ober light violet and a a s controlled by two coniplcmcntary genes.
I3trad;lr ct I I / (19971 reported the tnvolvement ill' tlircc gcncs lor col>\
pigmentation (PIC. PI. PC) and seed coat pigmentation ( P f , PI. P.1 and L>or gcncs lor pod
t ip pigmentation (Ptt, I'/l '?l'j) and flower colour (Pt P i P? P j ) i t1 the crosses tlicy
htudied.
tiuniar (1 097) reported cot i ip lc t i i t .~ i ta t i~~~i Ibr pink Ilo\rer colour in two crosses ( i f
\ \ l ~ ~ t e Ilri\rereil clilchpca accc~sions. In the cross l'OO7.3 s RS I I . I:I bras pink and In 1::
tlo\\er coliiur scgrcgetcd In tlic rzrtio oI"J pitih : 7 \vliitc s l i oa~np coniplc~iicntary typc o f
gene nctlon.
Vijayalakslini~ (IOOX) studied the ililicritnlicc 01' Ilo\\er colour ;itid rcportcd Llircc
d i lk ret i t geties governtng I loucr c o l ~ ~ u r . and \upplcnicntnry typc 01' gcnc . ~ c t i ~ i n uab
ohcer\ed it1 tlie L\\o crosses studled and designated genotype Ihr ~ r l i i t c I h ~ a c r colour ic
( '( 'hhl'l', l'or pitik ( '( ' f lhP/~ and h r hluc ( '( '/1/1/1/1.
Kurnar c r a / . (2000) reported supplementary t!pe 11I'ge1ic action l i ~ r Ilo\ver colour
hascd on scgrcgatlon l i l r two independent loci hy crossing \%hire llower coloured lbm:~lc
p:irents u i t h blue t louercd mole parent.
Sabaghpour (2000) monogcnic inhcrltancc obtained fi>r the character. pink v \
\vhtte tlo\vers. and determined the genotype of ICCV 2 as I'l'hh(Y' and o f J ( i 02 as
PPRBC'C. I l e also suggested that the seed coat colour is controlled by Lhrcc pair?, orgcncs
but some of them were different than for flower colour.
(iaur and tiour (2001 ) reported that a rcccssnc gcnc. dcs~gnntcd rf~,. \\;IS I'bund to
inhihit I louer colour jvithout affecting \eil i colour o f corolla. When I/L. present in
honiogygous condition. 1'-B- and111,B- genot)pcs ga\c pink-tcincd \%Ii i~c Iluncr ,111d hlue
\cincd \rhitc Ilo\rcrs. rcspcct~vcl>
IFroni the aho\c stud~cs. !he ~ l i l l c r ~ ~ o ~ i c c o f Ilo\rcr c o l o ~ ~ r is reported lo he
riioliogcnlc. digcnic and trigcnic. I3irnda1. c.1 '11. (1007) proposed 4-gclie inlicritancc
model kir Ilo\rcr colour. I llus, a1 Icost lhrcc gene\ arc go\crnlllg 11ic Ilo\rcr colour in
cliichpcu, frigenic inlicritnncc model %;I\ propmcd hy 11));lr :111d I~:I~~IsLI~~:II~~~I~I;II~
I 1430). 1)'C'rur and I c~idulhar ( 1')70) and I ' l iad~i~s (1076) I Iic gc~ic symhols ( ' M and I'
gi\cn h) Ayyar and I3alasuhra1iian1i11i (10.30) ;111d /'CO(~. I'L.o~I il~ id I'co/,) gi\ 'c~i h>
1)'Cruc. and 'frndulkar (1970). Viiayolaksh~iii ( Ic)OX) :111d (iilur illid (;our (2001) could hc
Yanlc. as no allelic tests have becn conducted.
Iligcnii. inhcrilance modcl was proposed by Khan ;lnd Aklitz~r (1034). I':il (1934).
Ki~dnni c f (11. (1941). I'atil and Dcshmukli (1075). Reddy and C'liopde (1077). I';~\\ar and
I'atil (1070). Rao et (11. (Ic)80) and Kaln~ e / ul. (1004). I)avis ( IOOl) and Kurnar (1097)
I he different gene dcsiynatil)n> given hy thcse ~ o r k c r s ~iamcly l j and 1'. flco and .Sco,
I.YL,II and IVco and I'co, and IJcoh could b t same.
Al l the other workers have proposed monl)genic inhe1itani.e model. The gene
symbols used by them namely BCO, P, PCO, Pkco illid LIYO could he probably tlic same.
Thus, the segregation for one, two or three gene pairs \rill depend on the genetic
cc;nstitution of the parents. Some workers reported that genes for corolla colour had
pleiotropic effect on seed coat colour and seed sllapc while some others reported linkage
between genes for floirer colour. seed coat colour and seed shape.
It is apparent that the flower colour in chickpea 1s controlled hy Inore than tllrcc
gene pairs. The seed coat colour is also govcrncd b) more than three genes. At least one
gene for flower and seed coat colourq is common. As \re conduct more research more
gcncs will he identilied. 'l'liis is expected hecause the related specie? \rith \rhich chickpea
ahares considerable synteny l'i~tmt .scr/iv~ini, has many more genes identified for this and
other traits.
2.3 Correlation coefficients
Correlation coefficient is an important statistical tool fcir detennln~ng association
between t\ro characters. Ilnderstanding (if thc inter-relationship hetireen sccd )ield and
its components and among the components themsclvea is necessary to impro~e seed y~cld
since it is a complex character A review of literature for correlations of yield \\it11 yield
contributing traits is given as follo\rs
Redd) and Rao (1988) anal)zed 50 F2 chichpcn populal~ons der~icd from Inter
varietal crosses and rcported that seed and pod number per plant were positiiel)
associated uith yield per plant. Plant height had significant positi~e association with 100-
seed ueight. Number of. pods per plant was positively associated with numhcr ol.sceds
per plant. Plant height and 100-seed weight showed nnn-significant assoc~ation with
yield.
Sharrna and Maloo (1988) observed in 21 chickpea iarieties that the grain yield
was significantly correlated hith number of pods per plant for two planting dates (i.e. 28
Noiembcr and 14 December) (r = 0.7 and r = 0.7. respectively), aith the number uf
primary branches per plant (r = 0.5) and 100-sccd weight (r= 0.7) fbr earlier planting
dates. Thcy also reported that days to flowering showed strong posititc correlation hith
days to maturity, and days to maturity cshihitcd negatitc and signiticant correlation aith
100-grain weight in case of second sowing at but11 genotypic 2nd phenotypic Icvels.
Salilnath and Bahl (1986). Sar~dhu el ol. (1988). hfishra rr oi. (IOXX). Sing11 c l ol.
(1989). Sandhu ei (11. (1989). Sandhu and Mandal (1989) and Tagore and Sing11 (1990)
carried out association studies and reported significant positive association of seed yield
ai th primary branches per plant. secondary branches per plant and pods per plant and
suggested selection tbr these characters to i ~ ~ ~ p r o i c yield.
Sandhu and Mandal (1989) Troni their studies in 48 ditcrsc chickpea lines
obseried that seed yield was posititcly correlated aith primary and sccondary hranches.
pod number and seed number.
Ali (1')90) conducted studies on six adianced lines ol'dcsi chickpea. compared
with two check cultiiars and found positive association of gram yreld with plant height
and grain mass. Iie suggested to consider longer durat~on of tlo\vcring, late nlaturity and
large grain mass white selecting genotypes for grain yield.
Uddin el ul. (1990) investigated correlation derived from the data of 54
genetically diverse chickpea lines and reported that yield per plant had significant
positive correlations with pods per plant. 100-seed weight and primary hrancheh per
plant.
Yadav (1000) conducted studies on F? popul;ltion of three cliichpea crosse.: ~ I i i c h
indicated that seed yield Has s~gnliicantly and positively correlated with numhcr of seeds
per plant, number of pods per plant, numbcr of hccondary hranches. 100-seed \+eight and
plant height.
Ilhambotha el (11. (1994) evaluated 32 genotypes in four difIicrcnt cnvlronments,
and their correlation analysis rexealed positive associat~ons hctween pod bcaring
branches per plant. pods per plant and plant height with seed yicld. Days to maturity had
a nun-significant correlation with yield in all four environnicnts.
Escr et ul. (1991) studied three varieties. ~ h i c h were calibrated according to sccd
size into large, medium and small types. 'l'hcy concluded that thc highest \'slues of yicld
and yield components were obtaincd [rom large seeds indicating the positive influence of
seed Inass on yield and its components.
Pundir et ul. (1991) found that lsaf and leaflet arca, and 100-seed ~ c i g h t were
positively correlated with each other but negati\'cly correlated nith secd protein content
ill 25 f'ic,c,r urietinum accessions. They also found negatlve correlation b e t ~ e e n 100-seed
weight and seeds per pod.
Abdali (1992) worked out correlations on I:4 and 1:3 generations of threc chickpea
crosses which revealed that grain yield expressed highly significant positive association
with number of pods (0.78-0.94). nulmber of seeds (0.79-0.93) and secondary hranches
(0.51-0.87) in all crosses in two generations. Number of pods per plant was significantly
and positively correlated with number of seeds per plant and secondary branches in three
crosses in both generations.
Akdag and Sehirali (1992) found significant positive correlations het\%een sced
yield per unit area and protein yield per unit area, and between seed yield pcr plant and
plant height, number of primary branches, pods and seeds per plant. 'They reporlcd
significant negative correlation between seed yield per plant and plant density.
Uouslama el crl. (1992) and Varghcse ct crl. (1993) reported s~gnificant positice
association of seed yield with pods per plant and 100-seed wigh t , and consldcrcd these
traits as Important yield components in selection of better gcnotypeh in chickpea.
Dasgupta o (11. (1992) ohserved significant and posi~ivc correlations of sccd yield with
pods per plant, seeds per plant and 100-seed weight. 'They obhcrved significant posit~ve
correlations betwecn seeds per plant and seeds per pod, and betwecn pod5 per plant and
seeds per plant in 28 genotypes of chickpea, They observed significant negative
correlation between seeds per pod and 100-seed weight.
Jadhav et 01. (1992) studied yield correlat~ons of irrigated ('icer urierin~~m and
safflower under various intercropping combinations. 'They found the number of
~roduct ive pods per plant and seeds per plant to be most highly correlated with seed yield
per plant, followed by number of total pods per plant. 1000-seed weight and number of'
branches per plant in C'iccr orirrinum.
Chavan ct ul. (1994) in a field study on I I chickpea cultivars grown under rainfed
and irrigated conditions found significant positive correlation between seed yield and
protein content and observed that irrigatio~~ significantly increased seed yicld and proteln
content.
Lal er (11. (1993) reported in chickpen genotypes that sccd yield was positively and
significantlj correlated with pod number and plant hcight. and negiltivelj correlated with
100-seed weight. I'lant licight showed significant negativc correlation w ~ t h branch
number. I'od number had significant negative correlat~on with 100-sccd wcight. fhcy
suggested pod numbcr and plant hcight as imporknnt charactcrs for seed yicld.
Salhc ct ul. (1993) studied six cultivars of chickpea and ~iotcd significant and
positive correlations of grain yield with number of grains pcr plant, 100-seed w i g h t and
number of filled pods per plant. Correlations were s~gnificant and positive between plant
height and 100-seed weight, number of branchcs and total number of pods per plant,
number of filled pods and number of grdins per plant and 100-seed weight. and numbcr
of grains per plant and number of seeds per pod. Negal~ve correlation was found between
100-seed weight and number of seeds per pod.
Arora and Kumar (1994) evaluated 40 Ciccr oric/inlim genotypes and observed
that seed yield per plant was positive11 associated with pods per plant, plant height and
weight from the data on 10 yield and growth characters in varicty I>(; 5 .
Sarvaliya and Goyal (1994) studied 76 chickpea genotlpes and revealed
significant association hetween yield and 100 sccd weight. plant height. t~umher of
primary branches per plant, number of pods per plant, days to maturity and days to
flowering at genotypic and phenotypic Icvcls.
Rao er 01. (1994) studied 44 varieties of (' urierinum and reported that seed yield
was positivcly correlated with primary hranches. secondaty branches, 100-seed weiglit
and pods per plant.
Singh and Rheenen (1994) crossed JO 62 and MS 24 and evaluated along with
their F1 and F2 a id backcross progenies and observcd that number of seeds per pod \bas
positively correlated with seed yield in segregating generations (r =0.18).
Bhattacharya e/ ul. (1995) studied the association u f yield and yield components
under soil moisture stress and non-stress environmental in chickpea by taking twelve
genetically different chickpea genotypes and reported that. under non-stress conditions,
seed yield is mainly influenced through extent of biological yield followed by effective
node number per plant and number of seeds per plant, white under stress conditions
maximum association was observed for biological yield lbllowed by plant height. Iiarvest
index and days to 50 % flowering.
Khorgade el irl. (1993) based on yield correlation dcrivrd tionl data on nine
characters in 30 chickpea genotypes grown under normal and late so\cn conditions.
reported that seed yield showed pos i t~~ ,e significant association with pods pcr plant.
hranches pcr plant and 100-seed weight. whrreas secds per pod harl sigliilicant negatii~c
association with seed yield per plant under both conditions.
Sandhu and Ivlangath (1095) htudicd 32 diverse genotype5 of chickpea and Sound
yield pcr plant showed sigtlificalit positivc association with primary hranches. pods per
plant and harvest index and negati1.e associations ai th plant height (45 days allcr sowing)
and days to flowering.
Mathur and Mathur (1996) showed significant positive corrclatlon of grain yicld
per plant with pods per plan and 100 grain \veight but negative correlation with plant
height in 34 chickpea varieties.
Ozdemir (1996) reported that number of pods per plant had a sign~ficant and
positi\e correlation with seed yield. although it had a negatlve direct ci'fecl. its Indirect
effect was positive via seed number and seed yield per plant.
Manlare er ul (1997) reported on 22 genotypes of c h ~ k p c a that grain 11eld pcr
plant had p o s ~ t ~ v e c o r r e l a t ~ o ~ ~ s s l t h number o t pods per plant, numher o t branches per
plant. 100-seed rvclght and nunlber o t grams per pod However, only numbcr o t pods per
plant e x h ~ b ~ t e d slgn~ticant correlatlon
('hand and Slngh (1997) studled 49 genotypes and ohaervcd that gram y e l d per
plant had s~gnlficant pos~tlvt. ~ o r r c l a t ~ o n u ~ t h riurnher of podv per plant and seed? per
plant It rvas observed that number o t pod\ pcr plant and seeds per plant are tlic most
y ~ e l d c o n t r ~ b u t ~ n g ~haracters In ehlckped
Vya?aldkshnil (1998) \ tud~cd t w crosses o t ch~ckpca and rcported pos~t lvc
correlatlon w ~ t h number of pods and secdb pcr plant, nuntber o t prlnlary and secondary
branches per plant and seeds pcr plant also ~ n t l u e n ~ c d the sccd yield d~rectly or
lndlrectly
Or el ul (1999) studled the phenotyp~c correlat~on.\ h e t ~ e c n d q s to first floucr.
pod numher and mean grain a c ~ g l i t among F2 populat~on\ dcr~ved trum c r o w s hetween
early t l u \ v c r ~ n ~ (dest) x l ~ t e f l o u , c r ~ n ~ (kabull) ~ u l t ~ v d r s and rcvedlcd n strong asso~la t lon
In the characters studled
Sahaghpour (2000) reported that the number of secondary branches per
plant. number of pods per plant, sced y ~ e l d per plant and number of sceds per plant
exh~hl ted hlgh correlated response to sclect~on w ~ t h yield per plot
Raghu (2001) observed that yield showed h ~ g h positive association with days to
first flowering and a negative association will1 days to 5 0 % flowering. days to maturity
in F7 generation of chickpea. In FZ generation of the cross. yield per plant cxliihited a
non-significant correlation with all (he parameters under study excepting thc ]lode
number, wherein yield sho\ved positive association with nodc number.
Prom the above rekiew, allnost all cases numhcr of pods per plant. nuniber of
primary and secondary branches showed positive correlation with seed yield. Correlation
of seed yield with plant Ileight. 100 seed weight and seeds per pod were positive in somc
studies while ncgatite in other studies. It is easier to mcasure yicld than to mcasurc the
number of pods, and seconday branches. It could be said that a visual nieasuremenl of
pod number could be morc cfficicnt.
CHAPTER 111
MATERIALS AND METHODS
The present inbesligation was undertaken to study the inheritance of stem
pigmentatio~i and flower colour. and to determine the genotypes of dirkrent sepregants
with ahi te flower colour and to determine the association among important tralts in
chickpea.
The experiment was conducted during the post-rainy season 2000-01 at the
International Crops Research Institute for the Semi-Arid 'l'ropics (ICRISAT) Patanchcru.
A.P. It is located at an altitude of 545 m above mean sea Icvel. latitude 17"3?' N and
longitude 78'16' E. The heather data during the crop gro\+.th period is giben in
Appendix 5 . '1 he research material was provided by the Chickpea GREP Unit ICKISAT.
The details of the materials and methods followed in this expcrlmcnt are fbrnishcd
hereunder.
3.1 MATERIALS
The experiment was conducted ai th six different genotypes inbulvcd in three
crosses. The parents uscd were: ICC' 5716. ICC 17101. T-I-A. T 39-A. ICCV 2, and
RS 11.
During 1999. a chickpea accession ICC 5716 with highly purple stem and p~nk
flower colour was crossed to ICC 17101 with low stem pigmentation and ahi te flower
colour. The resulting F2 produced plants of four types I e , high pigmentation with white
flower colour, high pigmentation with pink flower colour, low pigmentation with white
flower colour and low pigmentation with pink flower colour. These four phenotypes were
planted to investigate the inheritance of stem pigmentation and flower colour. For the
stem pigmentation and floaer colour studies. both parents I e . . ICC 5716 and ICC 17101
and 151 Fl plant seeds were used for planting.
The remaining four genotypes were used to determine the different genotypes for
white flowcr. ICCV 2 and RS 11 have white t loacr colour and I-I-A and r 39-1 are
blue coloured types. Crosses were made between (ICCV 2 x T-I-A) and (RS 1 I x 'I' 39-1)
during 1997 rabr season in the glass house. The two resulting I:, 's produced plnk iloaer
colour. The F2 generations of these crosses exhibited three types of floaer colour; pink.
white and blue. The white flowcr coloured plants from the F2 of lCCV 2 1 T-1-A were
crossed a i th the white coloured 1:2 plants of RS I I \ T 39-1 in the 1998 nlh, season. And
one reciprocal cross was made. The FI of these crosses resulted in plnk and blue tlowers
and in F2 pink blue and white were resulted. From these three types only white flower
colour plant seeds were used as planting material in the 2000101 m h , heason. a total of
146 white flower plant seeds, 4 parents and 6 selfed white flowcr seed of prevlous cross
were used as planling material.
3.2 METHODS
The above mentioned chickpea plantlng rnater~als of !\ra sub experiments were
sown during the post rainy season (rahi) 2000-01 at ICRISA'T research farm Patancheru
on 20 October, 2000 on deep vertisol type soils under conserved soil moisture conditions.
The plots were single row 2 m long with 60 cm between rows. The seed to secd spacing
was 10 cm. 'Shere were 156 plots. Alpha design was used for these experiments with two
replications. In both the cases soaing was done with the help of planter. All necessary
cultural operations and plant protection measures were done to raise a healthy crop.
3.3 COLLECTION O F DATA
In stem pigmentation and tlower colour study, hoth ytem pigmentation and Ilo\ver
colour and in the determination of the of diffcrcnt genotypes fur ahi te flower colour
study the flower colour was recorded after full blooming for cach plot. Days to iirst
flower. days to first pod. days to 50 % flowering and days to ~naturity were recorded on
the average pcrformancc of plot basis. Single plant data on yield and yield contributing
characters were also taken from those selected plants in both cases.
3.3.1 Initial plant stand (IPS)
The total number ofplants at germination (about I0 days after soaing).
3.3.2 Flower colour (FC)
Colour of [he freshly opened tloaer i.e.. pink. bluc or whitc has recorded tbr each
plant
3.3.3 Stem Pigmentation (SP)
The stem pigmentation was observed at 25 days after sowing as high, low or no
pigmentation.
3.3.4 Plant height (Ht)
Height of the plant in centimeters was measured and recorded from the soil
surface to the top of the longest branch at the time of maturit> Ibr each sclccted plant.
3.3.5 Plant width (Wd)
Top canopy width in centimeters was riieasured and recorded for cach sclcctcd
plant.
3.3.6 Number of primary branches per plant (Ph)
At maturity, number of branches or~ginatlrig l i o ~ n the hase of the plant was
counted and recorded for each aelected plant.
3.3.7 Number of secondary branches per plant (Sh)
At maturity, the number of branchcs originating from all primary branches was
counted and recordcd
3.3.8 Numher of pods per plant (NPP)
Total number of pods both filled and unfilled, on all branches of' a plant was
counted and recorded.
3.3.9 Number of seeds per plant (NSP)
Total number of seeds obtained after threshing of all pods of a plant was counted
and recorded. 111 filled seeds and broken seeds were rejected.
3.3.10 Weight of 100 seed (HSW)
Weight of 100 seed was recorded in grams and was obtained b) the formula
Seed yield pcr p l ~ n t (g) - - - - - - - - - - - - -. . - -. - - - - - - - - - - - - - - - - -. . . - -. - - x 100 Total number of seeds per plant
3.3.11 Seed yield per plant (Yld-P)
All the seeds from each plant were ueighed Lrith thc help of Mettler's electron~c
wcighing niachine and recorded in grams.
3.4 STATISTICAL ANALYSIS
She data recorded on various characters studled uerc subjected to the Ibllowing
statistical analysis.
3.4.1 X2 test of good ness of fit
This is a test of statistical significance that is uscd to tcst the significance of difference
bctween observed and expectcd \.slues and also to test the val~dity of segregation rated
for detection of liukage and study of genctics. The %' tcst was also used to test thc
presence of linkage.
3.4.2 Correlation coefficients
Correlation rcfers to the degrcc and direction of nssociat~on between two or more
than two variables. Correlation coefficient measurc the mutual relationship bctween
various plant characters and determines the component charactcrs on which selection can
be based for genetic improvement of yield.
Simple correlation coefficient anlong yield and yield contrihuting traits were worked out
using the formula suggested by Panse and Sukhal~ne (1967)
(I sy Correlation coefficient (r) = -------------------
o x . n y
where; - -
Z f (x-x) (y-y) CTdx-dy ox) '= ---------.----.---. = . . . . - - - - - -. - -. . -. . - - -.
N N
o xy = mean product movement (or) the covariarico herween x atid y
o x = standard deviation of x
(I y = standard deviation of y
d x Zdy = dcviations
Significance of the correlation coefficient
r is estimated from 'n' pairs. l'he siglnficancc of corrciatinns was tested hy rcfcrring to
standard 't' table givcn by Snedecor and Cochran (1968) at 5 % and I % levels of
significance
Table 1 : Characters of parents used for determination of different white flower Genotypes
Character i ICCV2 RS 11
, ccBBPf
White
Semi erect
Desi
Angular
Medium
Medium
Genotype
Flower colour
Growth habit
Seed type
1 Seed shape
Duration
Pod size
h h
White
Erect
Kabuli
Owl's hcad
Very short
Large
T 3 9 - 1
CYC'BBpp
Blue
Sen11 erect
i n t e r a t e
r l C'C'BBpp 1 Blue
Spread
lntermediate--
Pea
Long
CHAPTER IV
RESULTS
During the post-rainy season of 2000-01 an experiment was conducted at
ICRISA'I Patancheru, near Hyderabad. Andhra Pradesh, ro study Ihe ~nheritancc
of stem pigmentation and f l o ~ c r colour. and to determine the association among
the important traits. Data were recorded on days to first I lo~er ing (DFF), days to
first pod formation (DFP), days to 50% flowering (DF50%), days to maturity
(DM), flower colour (FC), stcm pigmentation (SP). Plant height (I 11). plant width
(Wd). number of primary branches per plant (Ph), number ol'sccondary brariches
per plant (Sh), number of pods per plant (NI'I'). number of sccds per plant (NSP).
yield per plant (Yld-P). and 100-sccd weight (IISW). The results arc presented
under the following headings'
4.1 Inheritance of stem pigmentation and Iloaer colour
4.3 Iletermination of genotypes for white flower colour
4.3 ilssociations among important traits
4.1 Inheritance of stem pigmentation and flower colour
4.1.1 Stem pigmentation
The inheritance of stcm pigmentation (anthocjmin) was studied in the 1:2 and FI
generations of the cross ICC 5716 x ICC 17101. In the F2generation 106 low and 48 high
pigmented plants were obtained, Based on one-gene segregation and ~ i t h lo\+
3 3 pigmentation dominant to high pigmentation, Illis fits well to the expectcd ratio of 3:l
(Table 2).
The data for F3 generation of the above cross. confirmed the results of
F2 generation indicating that the stem pigmentation is controlled by a single gene
(Table 3).
1.1.2Flower colour
In this cross this cross thc inheritance of tlower colour was also studied, Tllc
ohservations obtained in F? generation ac re 121 pink: 33 ahite. Based on one geilc
segregation these fit s e l l to the expected ratio of3:I ('l'ahle 2).
Thc F3 generation of the above cross also confirmed the results of b'2 generation
segregation, indicating that the pink Iloher colour is controlled by a singlc gene
(Table 3).
Table 2: Segregation for stem pigmentation and flower colour in the FZ generation of the cross ICC 5716 x ICC 17101
I Character i Phenotype
FhJaer colour
I
LON
lligh
Pink
White
--
Plafc I i':~t.cnt$ showing high pignlentcd stem ICC5716,los pigmcnled 4tcm ICC 17101
IJiate 2 IJarentu sho\\ing pink tlo!~crlC'('5716 and nhite ilower I('C'1710J
Table 3: Segregation for stem pigmentation and flower colour in the F3 generation of the cross ICC 5716 x ICC 17101
Cross Character
ICC 5716 x Stem ICC 17101 pigmentation (F3 generation)
Flower colour
Pheno Observed Number
Lou-
Pink 683
White 277
Ap~r0pri.t. 1 1 p 1 Ratio
4.1.3 Joint segregation of stem pigmentation and flower colour
In the F2 population the joint segregation of stem pigmentation and tlower colour
was studied together. Four classes of stem pigmentation and flower colour were obtaincd
indicating that two genes governed these traits (Table 4). The four classes; low
pigmentation with pink flower. low pigmentation with white flower, high pigmentation
with pink flower and high pigmentation with white flower fit well to a 9: 3: 3:t ratio
value was 4.30" at 3 d.t). F3 generation data confirmed the results of F1 generation
(Appendix no. I). This indicated that the two genes segregated independent of each other.
Table 4: Joint segregation of stem pigmentation and flower colour in F2 generation of the cross ICC 5716 x ICC 17101
Observed
number
Low pigmented with pink flowers
Low pigmented with white flowers
High pigmented with pink flowers
High pigmented with white flowers
Total
4.2 Determination of different genotypes of white flower colour
The crosses between ICCV 2 (white) x T-I-A (blue) and RS I l(uhite) x T 39-1
(blue) produced pink flower colour in F I indicating interaction of bluc with white flower
colour. In the F2 generation of both crosses were resulted in three typcs of flower colours
pink, blue and white indicating supplementary type of gene action and their expected
genotypes were showed. (Table 5 ) .
By crossing these white flowers ( F2 white flower of (ICCV 2 x T-I-A) x
F2 white flower of (RS 11 x T 39-1)) produced pink and bluc flowers .The F2 generation
of these cross resulted in three types of flower colours pink. blue and white. The expected
Table 6: Table showing possibility of crosses amongwhite flowers from the crosses of ICCV 2x T-1-A and RS 11 x T 39-1
Note: Selfing the high lighted phenotypes can produce (ripple recessive genotype for white flower colour
ICCV 2 x T-1-A
Fz white flower
genotypes
RS I1 x T 39-1 F1 white flower
C'ChhPP
1
CC'hhPp
2
(T'hbpp
1
ccBBPp
2
pink:bluc
?:?
pink: blue
3:l
pink:hlue
1:I
genotypes ccBBPP
I
pink
1
pink
2
pink
1
ccBBpp
1
pink
1
pink: blue
1 : l
blue
1
Table 7: Crossing pattern and segregation of white flower types from both crosses
(lCCV2 x T-l- F2 white ) x (RS I I x T 39- 1 F2 white)
. . for getting triple recessive genotype for white flower colour
Cross
Complim- cntary gene action
FI
F2
I I I I I
Gene- ration
F2 white ) x (US l l x T 3 9 - 1 F2 white
White x White
Pink
Pink White
I I I I
FI
F2
Pink
Phenotype 1 Expected 1 Genotype
(TbbPP x ccBBPP
CcBbPP
('-B-PP ('-hhPP ccB-PP
ICCV2 x T - l -
Plnk C7cBbPp 1 blue CcBbpp
Appropriate Ratio
White x
Supple- lnentary gene action
Type of interaction
C('hhPn x
P~nk Blue Wh~te
C.B-P- ( -B-pp (-hhP- C -hhpp ccB-P- cc B-pp tcbbP-
Complim- elltary gene action
F Blue
ICCV2xT-I-A F white ) x (RS l l x T 39-1 F2 white
Blue Wh~te
Complim- entary gene action
'-B-pp ('-hhpp ccB-pp cchhpp
F2 Blue ('6-pp White C-bbpp
ccB-pp cchbpp
.; n '1 J
4.3 Association among important traits in the cross
4.3.1 Associations among important traits in the cross ICC5716 x ICC17101
4.3.1.1 Days to 50 % flowering
Days to 50 % flowering was significant and positively correlated with days to
first flowering (0.693**), days to first pod formation (0.600**), days to maturity
(0.261**) and with I00 seed weight (0.198**) in F2 generation.
Days to 50 % flowering was not having any association with recorded ch;lracters
in F3 generation
4.3.1.2 Days to first flowering
Days to first flowering was significant and positively correlated with days to first
pod formation (0.921**), days to maturity (0.274**) and with 100 seed weight (0,172')
in F2 generation.
Days to first flowering was significant and positively correlated nith days to first
pod formation (0.803**) and significantly negative correlation with initial plant stand
(-0.308**) in F3 generation.
4.3.1.3 Days to first pod
Days first pod formation was significant and positively correlated with days to
maturity (0.230**), 100-seed weight (0.200*) and with days to 50 per cent flowering
(0.600**) in F2 generation.
Days first pod was significant and positively correlated with days first flowering
(0,083**) and negatively correlated with initial plant stand (-0.399**) in Fj generation.
4.3.1.4 Days to maturity
Days to maturity was significant and positively correlated with days to 50 per cent
flowering (0.2611*), days to first flowering (0.274") and days to first pod formation
(0.230") while negatively correlated significantly with plant height (-0.208**). initial
plant stand (-0.164') and with number of pods per plant (-0.169*) in F2 generation.
Days to maturity was not having any correlations with the recorded characters in
F3 generation
4.3.1.5 100 seed weight
Hundred seed weight significant and positively correlated with days to 50 %
flowering (0.198*) and with days to first tlowering (0.172*) in F2 generation.
Hundred seed weight was not having any association with the recorded characters
in F3 generation.
4.3.1.6 Plant height
Plant height wps significant and positively correlated with number of pods per
plant (0.458**), number of seeds per plant (0.455**), number of secondary branches
(0.375**), plant width (0.606**) and with yield per plant (0.500**) in F2 generation.
Plant height was significant and positively correlated with number of pods per
plant (0.435**), number of seeds per plant (0.455"). number of primary branches per
plant (0.319**), number of secondary branches (0.228**), plant width (0.483**) and with
yield per plant (0.547**) in F3 generation.
4.3.1.7 Initial plant stand
initial plant stand was significant and negatively correlated with days to maturity
(-0.164*) and with other characters no association is observed in F2 generation.
Initial plant stand was negatively correlated significantly with days to first
flowering (-0.308**) and with days to first pod formation (-0.399**) in Fj generation.
4.3.1.8 Number of pods per plant
Number of pods per plant was significant and positively correlated with plant
height (0.458**), number of seeds per plant (0.893**), number of secondary branches
per plant (0.395**), width of the plant (0.539**), yield per plant (0.864**) and was
negatively correlated significantly with days to maturity (-0.169*) in F2 generation.
Number of pods per plant was significant and positively correlated with plant
height (0.435**), width of the plant (0.358**), number of seeds per plant (0.802**),
number of primary branches per plant (0.275**), number of secondary branches pcr plant
(0.252**) and with yield per plant (0.813**) in F3 generation.
4.3.1.9 Number of seeds per plant
Number of seed per plant was significant and positively correlated with plant
height (0.455**), width of the plant (0.568**), numbcr of secondary branches (0.45 I**)
and with yield per plant (0.946**) in F2 generation.
Number of seed per plmt was significant and positively correlated with plant
height (0.455**), width of the plant (0.382**), nurnbcr of prinlary branches (0.275**),
number of secondary branches (0.262**) and with yield per plant (0.875**) in F3
generation.
4.3.1.10 Number of primary branches per plant
Number of primary branches per plant was not having any association with the
recorded characters in F2 generation.
Number of branches per plant was significant and positively correlated
with height of the plant (0.319**), width of the plant ().344**), number of secondary
branches per plant (0.304**) and with yield per plant (0.312**) in F3 generation.
4.3.1.11 Number of secondary branches per plant
Number of secondary branches per plant was significant and positively correlated
with plant height (0.375**), width of the plant (0.378**), number of pods per plant
(0.395**) and with number of seeds per plant (0.45 I**) in F2 generation.
Number of secondary branches per plant was significant and positively correlated
with plant height (0.228**), number of primary branches pcr plant (0.304**), number of
pods per plant (0.252**) and with number of seeds per plant (0.262**) in FI generation.
4.3.1.12 Width of the plant
Width of the plant \bas significant and positively correlated with plant height
(0.606**), number of pods per plant (0.530**), number of seeds per plant (0.568**) and
with number of secondary branches per plant (0.378*') in F2 generation.
Width of the plant was positively correlated significantly with plant height
(0.483**), number of pods per plant (0.358*'), number of seeds per plant (0.382") and
with number of primary branches per plant (0.344**) in FJ generation.
4.3.1.13 Yield per plant
Yield per plant was significant and positively correlated with plant height
(0.500**), width of the plant (0.566**), number of pods per plant (0.864**), number of
seeds per plant (0.946**), and with number of secondary branches Per plant (0.447**) in
E 2 generation.
1 J Yield per plant was significant and positively correlated with plant height (0.547**),
width of the plant (0.417'*), number of pods per plant (0.813**), number of seeds per
plant (0.875**), with number of primary branches per plant (0.312**) and with number
of seconday branches per plant (0.312**) in F3 generation.
4.3.2 Associations among different white flower colour genotypes
In the cross F2 white flower ICCV 2 x T-I-A x F2 white flower RS I I x T 39-1
the relationships of quantitative characters were studied among white flower genotypes
to find out the association among traits. Correlation coefficients were calculated. The
magnitude and direction of association among morphological traits in this cross were
presented below.
4.3.2.1 100 Seed weight:
Ilundred Seed Weight was correlated negatively significant with days to maturity
(-0.174') and with number of pods per plant (-0.286"' and with number of seeds per
plant (-0.182' ).
4.3.2.2 Days to 50% flowering
Days to 50% flowering was a correlated positively highly significant with days to
first flower (0.791") and with days to first pod (0.779") and with days to maturity
(0,409' ) and with plant height. (0.46") and with number of secondary branches (0.293")
and with plant width (0.392").
Table 4 b Correlation coefficients for the F2 generation of ICC5716 x ICC 17101
D505CF DFF DFP DM HSW HT IPS NPP NSP PB SB WD Yld-P
D50%F 1 DFF 0.693" 1
DFP 06" 0 921" 1 DM 0.261" 0.274" 0 23' 1
HSW 0.198' 0.172' 0 2 0.055 1
HT 0.016 -0 041 -0.085 0.208" 0.03 1 IPS -0 12 -0 061 -0 023 0 164' -0 144 -0.022 1 NPP -0.108 -0.098 -0.065 0 169' 0005 0458" 0 128 1 NSP -0.106 -0 086 -0.075 -0 111 -0 025 0.455" 0.063 0 893" 1 PB 0 054 -0 022 0 018 -0 065 0.149 0 121 0.079 -0 072 -0.097 1 SB 0.048 -0 105 -0.149 0 091 0 065 0.375" -0 104 0.395" 0.451" 0.132 1 WD 0.019 -0 027 -0.05 -0 002 -0 075 0 606" 0 043 0 539" 0.568" 0 146 0 378" 1
Yld-P -0.108 -0 087 -0 06 -0 116 0 109 0 5 0.07 0 864" 0.946" -0 056 0 447" 0 566 1
+ Significant at 5% level "S~gnlficant at I%ievel
4.3.2.3 Days to first flowering
Day to first tlower was posi!i\cl! corrclotcd higlil! ~igtltt ic;~m \ r ~ t h d;l!.s to first
pod (0.941") and with days to maturily (0.266") and \ t i ~ h plant licigll~ (O.J!(I") mid \tit11
number of secondary branches (0 338") and w ~ t h wldlh ol'lhc pl;itn ((1 35s")
4.3.2.4 Days to first pod
Days to lirst pod \\as pnsiti~cly corrclnlcd s ~ g n ~ l i i . : ~ ~ i ~ \$it11 d ,~ys [ I ) ~ii;iluril!
(0.297"), and with plant Ileigh! ((1.101"' ntld \ \ ~ l I i tirlmhcr 01' scco~lil;~ry lhr,inclics pcr
plants (0.310") and with ~ i d t h of the plant ((1.359").
4.3.2.5 Days to maturity
Days to maturity was posilivel! corrcliitcd ;ind signtlic;im \rill1 pl:~nl h c ~ g h ~
(0.312") and ~ i t h number of secondark hranchch (0 .170 ' ) ;ind uilh \rldtli 0 1 l l ~ c plotit
(0.334").
4.3.2.6 Plant hcight
I'lant he igh~ \\as correla~cd pcrsil~vcly highly r~gn~licaii! \rltti ~iuliihcr i~l 'piids per
plant (0.272") and with days to maturit! (0,312") and uitti uitlth ol'llic planl (11.011?"1
and also highl) signiticantl) correlated \ + ~ t h d a j s to .5O?C t loacr~ng (0 440") iind uith
days to first flowering (0.420") and \ritli days to lirst pot1 ibrmiit~on ((1 401") and \cilh
days to maturity (0.3 I?").
4.3.2.7 Number of pods per plant
Number of pods per plant aas positivel) correlated s~gn~l icant ly uith plant hc~ghl
(0.272") and with number of secondav branches per plant ((1.51 he ' ) and w ~ t h numhcr ol'
seeds per plant (0.418") and with width of the plant (0.428") and w ~ t h !~eld per plant
(0.205") but number of pods per plant was negativclj correlaled \vith 100 seed \\eight
(-0.286").
4.3.2.8 Plant width
Plant width correlated highl) significan~ \\ith days ro 50% I l o ~ \ e r i ~ ~ g ((l.jl)?'*) i111cl
with days to first flower (0.355") and wit11 daks lirsr pod (0.350") :111t1 \ \ i t11 ~ : I ) s 10
maturity (0.334") and with plant lheight (0.602") and \\irll I I \ I I I I ~ C ~ 01' pod^ per pIil111
(0.428").
4.3.2.9 Yield per plant
Yield per plant is not I~aving an) associa~ions \ r l t l~ lilllcr rcccirdctl c I I ; I ~ : I c I ~ ~ \ c\ccpt
with number of pods per plant (0.?5.5**)
4.3.2.10 Number of primary branches per plant
Number of primary branches per plan{ is !la1 h,lving on! o\cic!oll~lll~ \\!rh orher
the recorded characters which were s~udicd .
CHAPTER V
DISCUSSION
This study was conducted to investigate inherilancc of' sten1 ptgn~entalion and
flower colour. to determine different genotypes for white flower colour and to co t~~pu tc
associations among important traits in two crosses ol'chickpea. 'The results are discussed
under the following headings:
5.1 Inheritance of stem pigmentation and flower colour.
5.2 Determination of different genotypes for white flower colour.
5.3 Associations among important traits.
5.1 Inheritance of stem pigmentation and flower colour.
Flower colour has profbund cffect on other morphological and physiological
patterns. 'l'he inheritance of stem pigmentation and llowcr colour was studicd in chickpea
cross ICC 5716 x ICC 17101. ICC 5716 has light purple stem and pink Ilowcrs. ICC
17101 has high purple stem and white tlowcrs.
'The stem pigmentation was controlled by a single gcne pair in the present study.
Similar results were obtained by Bhapkar and I'atil (1962), Aher and Patil (1984). Singh
and Singh (1995). l'efera (1998) and Sahaghpour (2000). In the present study low stem
pigmentation was found to be dominant over htgh stem pigmentation. Ilo\vcver, 1:ssomba
er ul. (1987). Metz er a/. (1992). Singh er u/ (1993). Ghatge (1994a). Mathur (1998).
'refera (1998). Venugopal and Goud (1998) and Sahaghpour (2000) reported that high
pigmentation was dominant to lowlno pigmentation. Ghatge and Kolhe (1985). Kabir
and Sen (1991), Karkannavar ei 01. (1991). Ghatge (1993) (1994a) md Mathur (1098)
reported digenic control for the stem pigmentation. In addition to this complimentary
type of gene action was reported by Ghatge and Kolhe (1985). Karkannavar r i 111 (1901)
Mathur (1998) and Venugopal and Goud (1998) for pigmented and nollow pigmented
characters.
In chickpea purple foliage colour could be used as a marker in identification of
true hybrids (Sandhu r / oi., 1993). Mathur (1989) reported that this purple pigmentation
depends on sunlight. Sandhu rr ul. (1993) indicated that pigmentation remained stable
from seedling stage to plant maturity in ICC 6071.
Flower colour is also an imponant morphological marker in chickpea. In this
study pink flower colour is dominant over white flower color indicating that a single gene
governs this character. 'These results support those of I'impl~kar (1943). Khan ' 1 111
(1950). Bhapkar and Patil (1962 and 1963). Gil and Cuhero (l993), I'ct'era (1008) and
Sahaghpour (2000), suggesting that the ~ h i t e flower colour is recessive to thc pink
flower color. But Khan and Akthar (1934). Kadam ei oi . (1941) I'awar and I'atil (1079)
Ghatge (1994a) and Kumar (1997) reported that two genes controlled this character.
Ayyar and Balasubramanian (1936). D 'c ru~ and Tendulkar (1970). Phadnis (1976),
Vijayalakshnli (1998) and Kumar ei oi (2000) suggested trigcnic inheritilncc for this
character. According to them all the three genes I e C, B and I' should hc present in thc
dominant condition to produce pink flower colour. When ( ' and B are in dominant
condition blue colour is produced and when either B or ( ' is in homozygous recessive
condition white flower colour IS produced.
In the joint segregation for stem pigmentation and flowcr colour. these two
characters segregated in a 9:3:3:1 ratio indicating that both the characters were governed
by two different genes that showed independent assortment, These results vary froni
those proposed by Aher and Patil (1984). Ghatgc (1994b). Joshi el (11. (1994). 'Tcfcra
(1998), and Sabaghpour (2000). 'They found that the gene that governs the flower colour
has pleiotropic effect on stem pigmentation. From the above discussion stem
pigmentation and flower colour can show nionogenic, digenic and trigenic inheritance
depending on the parental genotypes.
5.2 Determination of different genotypes of white flower colaur:
In chickpea, three distinct tlower colours are identified namely pink. hluc and
white. The flower colour is an important trait since it is a reliable morphological marker
for comparing chickpea accessions. Most of the dcsi varieties of chickpea arc of pink
flowered type and kabuli types always have white flowers. White llowcr accc?sions
account for about one third of the world gcrmplasni and those with blue llowcrs arc rare
(Pundir et al., 1988).
The cross between two white flowered parents ICCV 2 and RS 11 produced pink
flower. These two parents when crossed to blue tlowcred parents 'r-I-A and 'r 39-1 also
produced pink flowers. This indicated an interaction between the genes for white and
blue flower colours resulting in the formation of pink flowers in the Fls. This also
suggested the involvement of more than one gene in governing the flower colour. These
results differ from those of Pimplikar (1943). Khan el a1 .(1950), Bhapkar and Patil
(1963). Patil (1964). Athwal and Brar (1967). Patil (1967). Khosh-Khui and Niknejad
(19711, More and D'cruz (1976). Nayecm el (11. (1977). Reddy and Nayeem ( I 978).
K u m a el u i . (19821, Pawar and Patil (1982 and 1983). Kidambi el 111. (1988) Singh er 111.
(1988). Gil and Cubero (1993) and Pundir and Reddy (1997) who propclsrd nionogcnic
inheritance model.
1.'2 of these crosses segregated in tile ratio of 9 pink: 3 blue: and 4 white flower
colour individuals. indicating the supplementary type of gene action and digcnic control
of this character. Iligenic inheritance model was proposed by Khan and Akhtar (1934).
Pal (1934). Kadam el ul (1941). More and D'cruz (1970). Patil and I)eshmukh (1975).
Reddy and Chopde (1977). Pawar and I'atil(1979). Rao el ol (1980). Davis (1991) and
Kumar (1997). According to the digenic model assuming gene designation are proposed
by Khan and Akhtar (1934). a dominant factor D produced blue colour. A Sactor P
produced pink colour in the presence of B but hy itself produced no colour In the
absence of B, the flowers were white. whether I' was presenl or not indicating the
epistatic actlon of hh.
The different gene symbols given by different scientists. namely B B ( ' 0 , I,Rc'O.
and PCO, for blue colour and P, ,Sco, W ( ' 0 and /'('Oh showing supplementary gene
action could represent the same loci as they werc designated without conducting the
allclic tests. The segregation ratio of 9 pink : 3 blue : 4 white flower colours in the Fz
generation of both the crosses was indicative of similar genetic const~tution of the two
white flowered parents ICCV ? and RS-11. However, this was not the case. as the two
white flowered parents produced a pink FI when crossed (Kumar 1997) indicating
different genetic constitutions for their white colours. llence the digenic model of
inheritance was not found to be appropriate (Kumu sr (11. 2000).
The possible white flower color genotypes of the ahovc crosses will he n~lIBPP.
ccBBPp, ccBBpp from ICCV2 x 'T-I-A cross . ('( 'hhPP. ('C'hhPp and ('( 'hhPP from
RS I I x T 39-lcross.
Prom the intercrossing of the above mentioned differen[ genotypes for white
flower colour from the two crosses, the F I was pink. Fl segregated pink and white. In
this case the type of interaction ohserved is complimentary (9:7). So the expected
genotypes of the parents involved in this cross wcre ( ' ( 'hhPI1 x rt13/31JP. 'fhc genotype
for pink flower is ('-B-P- and the white tlower genotypes are C'-hh1'-, icB-1'- and cchhP-
In other cross between whitc flowered genotypes, pink and blue flowered
plants wcre produced. So the expected genotypes of the parents in\,olvcd in this cross
werc ccBBpp x ('('hhPp pink and blue tlowercd plants werc with C'cnhl'p and
('cBhpp genotypes. Sslfing these plants produced some tripple rccessivc genotypes for
white flower colour.
From the other cross between the white flowered genotypes in I:, pink and blue
flowers were resulted. So the expected genotypes of the parcnts involved in this cross
were CChhPP x CcBhpp, the F I pink genotype is C-B-P- and blue gcnotype is ('-B-pp.
Fl segregation of pink colour gave all h e e flower colours pink C'-B-P-, hluc with C'-B-
pp and white with CCbbpp. ('chhPP, CchhPp, C'C'hhpp. ccBBPP, ccBBPp. ccBBpP,
ccBbPP, ccBbPp ccBbpp, cchhPP, cchhPp and n,hbpp llerc also we got tripple
recessive genotypes and blue flowers segregated into blue C'-8-pp and white with ('-
hhpp, ccB-pp, ccB-Pp and cchhpp
Thus from this study flower colour uas observed to be governed by three genes
and white flowered genotypes can have different constilution as proposed by Ayyar and
Balasubramanian (1936) Davis (1991 ), Vijayalakshmi (I99R). I'cfera (1998). Kunlar
et r r l . (2000). In this study various genetic constitutions for white flower colour including
one tripple recessive homozygous genotype were determined.
Considering the occurance of various shades of the colours ohscrved in this study
and 22 genes known for flower colour in the related genus Pislrm ~t is apparent that more
than three loci govern flower colour in chickpea Kumar (1097). 1:urthcr studies are
therefore, warranted to investigate evolution of this character.
5.3 Association of traits in the cross ICC 5716 x ICC 17101
An understanding of the nature of association of yield and yield contributing
characters are of great significance in proper planning of selection programmes and
genetic improvement of these characteristics. The associat~on are from t h ~ s study are
discussed here under.
I n the cross ICC 5716 x ICC 1 7101 in F? and FI generations seed yield per plant
is positively correlated with plant height (0.50**) (0.547*+) respectively. Dut this is
varies with Reddy and Rao (1988). Jahhar and Manne (1991). Al i (1990). Yadav (1090).
Bhambota er u l (1994). Akdag and Sehirali (1092). Snthc er rrl (1993). Arora and
Kumar (1994). Lal er n l (1993). Sandhu and Mangath (1995) who reported negative
association with yield and no association was rcported by Reddy and Ran (1988). Jahhar
and Manne (1 991) and Vijayalakshimi ( 1098).
Yield is positively correlatcd \\ith number ol' pods per plant in I:] and
Pigenerations (0.864"). (0.813**) respectively. (Salimath and Bahl. 1986; Mishra
el cil. 1988; Reddy and Rao, 1988; Yadav. 1990; Abdali. 1992: Chavan c/ rri.. 1994;
Ozdcmer. 1996; Manjare cr 01.. 1997; Vijayalakshmi. 1998).
Yield per plant is highly positively corrclatcd w ~ t h numher o f seeds pcr plant in
both F2 and I;? generation. (0.946") and (0.875**) respectibely. supporting Reddy and
Rao (1988), Sandhu and MandaI(1989). Yadab (1990). Akdag and Sel~irali (1002). Sathc
er u l (1993). Chand and Singh (1997) Vijayalakshmi (1998) and Sahayhpour (2000).
Yield per plant is positibely correlated with number o f secondary hranchcs In I.'*
(Sharma and Maloo (1988); Uddin r r u l (1990): Abdali (1992): Rao er ul (1994);
Khorgade er u l (1995) and Manjare r r u1(1997), Vijayalakshmi (1998). Sabaghpour
(2000) i n Fj no association was observed for secondary branches.
Yield per plant is positively correlated with number o f primary branches per plant
in Fj. ( Salimath and Bahl, 1986, Mishra er a1 1988; Chavan er a1.,1994; Man~are cr ol.,
1997: and Vijayalakshimi. 1998 ). In FZ generation number ofprimary branches per plant
is having no association with yield per plant. Yield per plant is positively correlated with
width of the plant in both FZ and Fi generations.
Yield per plant shows non significant correlation in both I:? and FI generations
with, days to 50% flowering. But Ali (1990). Sawaliya and Goyal (1094). Sandhu and
Mangath (1995). Qayyum el ul. (1997) and Kaghu (2001) reported association \*.ith yield
per plant.
In both F2 and Figenerations yield pcr plant is showing non s~gnificant correlation
with days to maturity (Bhambotha er rrl.1994; Kaghu. 2001). hut ncgativc association
was observed by Qayyum e/ 01, (1997). Yield per plant is slwwing no corrclation with
days to 50% flowering in both F, and FI generations. But Kahman and I'artli (IOXX) and
Kaghu (2001) reported negative association w ~ t h yield per plant.
Yield per plant showing no association in hoth 1:2 and 1:) generations with days to
first pod formation. This is differing from Qayyunl er ul (1997) and Raghu (2001).Yield
per plant showed non significant association w ~ t h 100 seed \*.eight and initial plant stand.
5.3.1 Association of traits among different genotypes o f white flower colour
Phenotypic corrc!ation studies \*.ere carried out to find out thc association^ among
important traits from different white flower colour genotypes..
Yield per plant among white flo\*.er genotypes showed a non signilicant
association with all the traits under study (Bhambotha er u l , 1994 and Raghu. 2001)
except with number of pods per plant ( Manjare er u l 1997; Vijayalakshmi. 1998; Or
et a1 . 1999; Sabaghpour, 2000 ). Number of pods per plant is correlating with number of
secondary branches per plant ( Bejiga ef a / , 1991 : Chhina el ( I / . , 1991: Abdali.1992:
Vijayalakshmi. 1998). Plant height showed positive association with nuniber of
secondary branches per plant (Choudhary and Mian. 1988: Vijayalakshmi. 1998) and
with number of pods per plant it varies from Vijayalakshmi (1998) who suggested
negative association. Number of pods per plant is showing negative association with 100
seed weight ( Lal ef ui.. 1993: and Vijayalakshmi,l998) \chile number of seeds per plant
also showing negative association with 100-seed weight (Sandhu and Mandal.lO80;
Pundir el a / , 1991; Dasgupta rf trl. 1992: Sathc r~ id 1903). Days to 50% t lo~cr ing is
more positively associated with days to maturity ( Arora and Jeena. 1999: and Rughu,
2001), days to first flowering is tnorc positively associated with days to maturity
( Sharma and Maloo. 1988: Raghu. 2001).
The study is useful as it s h o ~ e d that stem pigmentation and flower colour are
not pleiotropic traits as has always been reported in the literature. 'fliis indicates (hat
different mechanisms are operating for these characters. I:urther studies are needed to
determine the genes for these traits in chickpea.
Development of tripple recessive genotype for white tlower colour will simplify
allelic tests for flower colour in future chickpea studies. These genotypes will be
registered as genetic stocks.
CHAPTER VI
SUMMARY
The inheritance of stem pigmentation and tlowur colour and the association
among important traits in chickpca (('iccr urirtinlrm I..) were studied at the Internntionol
Crops Research institute for the Semi-Arid Tropics (ICRISA'f) IJatanchcru, ncnr
Hyderabad, A.P. during the 2000-01 post- rainy season. The li~llowing t a o expcri~nents
were conducted:
Experiment l
In this experiment the inheritance of stem pigmentation and Iloacr colour was
investigated. 'Two accessions ICC 5716 and ICC 17101 ae re used as parents. 1CC 5716
has sten1 with low purple pigmentation with pink tlowers and IC'C 17101 has stcm with
high purple pignientation ai th white floacrs. Data was recorded on stem pigmentation
and flower colour. The Iblloaing results were obtained:
Monogenic inheritance was confirmed Ihr the two ~norphological chi~racters low
v , high pigmentation of stem and pink and whitc llowcr colour. Joint segregation Ibr
these two characters fit ae l l to the digenic ratio of 9:3:3: 1 for the data for I:, generation.
indicating independent assortment of the two genes. 'I'hc traits showed no plciotropic
effect for stem pigmentation and floaer colour. Low pigmented stem is dominant to thc
highly pigmented stem and pink floaer is dominant to the whitc flower colour. 'The 1::
generation results were confirmed by those for the I:i generation.
Correlations estimated among quantitative characters and yield in comparison of
F: generation with F, generation, in both F: and F, generations yield per plant was
5 9
correlated positively with plant height and with number of seeds per p l a t . number of'
secondary branches per plant and with width of the plants. In I:? number of primary
branches per plant, and in F1 days to maturity, 100 seed weight arc not having
associations with any other characters studied. In both and I:\ generations, nunlhcr of
pods per plant is positively corrclated with plant hcight, plant width, nulnbcr o f seeds per
plant. number of secondary hranches per plant and with yield per plant.
Number of seeds per plant in both F? and FI gcncrations, is positively corrclated
with plant height, number of secondary branches, plant width and with yield per plant.
100-seed weight in F2 generation positively correlated with days to 50% flowering and
with days to first flowering but in the F3 gencratlon no correlation is observed.
Experiment 11
This was conducted to determinc different genotypes for white llower coltrur. I:or
this, four parents were used. ICCV 2. RS I I , 1' 39-1, and 1'-]-A. ICCV 2 and RS. I I arc
white flowered and the other two are hlue flowered accessions. Crossing of' ICCV 2 x
T-I-A, and RS I I x T 39-1 produced pink flowers in the 1 1 generation. I:I gcncralion 01'
these crosses segregated into three types of flowers pink, blue and white in 9:3:4 ratio
indicating involvemenl of three gcnes and supplementary lype of gene action. 'l'hesc arc
probably C: B and P loci as earlier reported in the literature. 'Therefore, the genetic
constitution for the four parents and their respective [:I and F2 generations are as follows:
ICCV 2 (white) CC'hhPP, T-I-A (blue) C'CBBpp, F , CC'BhPP (pink) Fz ('-B-P- (pink),
C-B-pp (blue) and white C-hhP- and C-h-pp. RS 11 (white) ccBBPP, 'f 30-1 (blue)
CCBBPP. FI CcBBPp, F2 I-BBP- (pink), ('-BBpp (blue) a d fix white c,c,BB['. and
ccBBpp.
The white flowers from the two crosses were intercrossed which resulted in pink
and blue flowers. The FI generation of these crosses some pink llowercd plants
segregated into all three types; pink, blue and white showing (9:3:4) supplementary type
gene action, some blue tlowered plants segregated into blue and white tlowers and
indicating (9:7) complimentary type of action. I:roni both pink and bluc flowered plants
white tlowered progenies resulted that may have the tripple recessive genotypes.
Some white flower plant genetic constitution may be heterorygous condition. Ilowcver.
for the first time a chickpea with a tripple recessive ccbbpp genotype for white llowcr
colour has been determined.
Yield per plant among different white flower types showed no association with
any traits under study except with number of pods per plant. 100 seed weight was
negatively correlated with days to maturity and with numbcr of pods per plant with
number of seeds per plant. Days to 50% tlowering was positively higl~ly corrclatcd
significantly with days to first flowering, days to first pod, days to maturity, plant height
and plant width. Number of pods per plant was positively correlated significantly with
plant height, number of secondary branches, and with number of seeds per plant, and
with plant width and with yield per plant but positively correlated with 100 seed weight.
The study is useful as it showed that stem pigmentation aid llower colour are not
pleiotropic traits us has always been reported in the literature. Ue\.elopment of tripple
recessive genotype for white flower colvur will simplify allelic tests for llower colour in
future chickpea studies.
LITERATURE CITED
'Ahdali Q N 1992 Variation in somc agronomic chxacteristics in three populations of' chickpea (Cicer arierintim I,.). M.Sc. Thesis. Jordan University. Jordati.
Aher R P, Patil J A 1984 Inheritance studies on stem and seed coat colour in bengalgram. Journal of Mahwashtra Agricultural llniversities 9(2) : I5 I - 152.
Ahmed M and 'ranki M 1 1992 Inheritance of anthocyanin pigmentation in chilli (Capsrctim annltnl I,.). Capsicum Newsletter I I : 20-21.
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* : Original not seen
The patlern of "Literature cited" presented ahove is in accordance %it11 thc "(i~~idelincc" Il ir lllc?is presentation for Acl~arya N. G. Ka~lga Agricullural University, I lyderahnd.
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M d l d d l M d H d d H d d l S 1 1 M d l d d l M d H d d H d d l S L L M d l d d l . . d d l P 1 L M d l d d l . . d d 1 V L L . d d l . d d H d d l C l 1 . d d l . d dH d d l t l l
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dl d d l M ~ H d d ~ d d i p 0 1 ~ d i d d i . d d ~ d d i v 0 1
, d d l . . d d l 901 . d d l . . d d l EOL
M d l d d l . . d d l Z O L M d l d d l . . d d l Z O L M d l . M d H d d H d d l 1 0 1 M d l d d l M ~ H d d H d d lLOL . . M ~ H d d ~ d d ~ 001 . . M d ~ d d ~ d d u 001
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M d l d d l M d H d d H d d l L 6 M d l d d l . d d H d d l 1 6 . . . d d H d d H S 6 . . . d dH d dH 96
M d ? d o 1 M d H d d H d d l 5 6 M d l d d l M d H d d ~ d d l 5 6
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APPENDIX 2 lndlvldual plant dab of FZ genention of the cmsr ICC 6718 x ICC 17101
PLOT IPS DFF DSOW DFP DM HT WD PB SB NPP Yld-P NSP HSW 1 7 50 52 57 105 38 33 2 6 101 1 7 4 152 1144 1 7 50 52 57 107 32 29 3 4 140 204 162 1802 2 10 48 51 58 103 37 29 3 4 105 139 118 1177 2 10 48 51 56 106 27 24 3 6 68 128 162 12 59 2 10 48 51 56 106 30 26 2 4 102 156 132 11 81 2 10 48 51 56 106 28 29 2 1 95 124 105 1 1 8 2 10 48 51 56 106 34 35 2 4 121 156 136 11 47
3 11 49 51 58 106 30 40 4 2 90 112 105 1066 3 11 49 51 58 104 34 33 3 7 101 1 4 8 126 1174 3 11 49 51 58 105 34 24 3 2 82 1 2 5 102 1225 3 11 49 51 58 104 43 40 1 4 203 249 203 1226
3 11 49 51 58 104 38 38 5 2 124 1 8 1 145 1248
3 11 49 51 58 104 38 29 2 3 134 17 9 156 11 47 4 11 41 48 49 102 33 34 4 2 128 191 153 1248 4 11 41 48 49 104 41 30 3 8 102 157 124 1268
4 11 41 49 48 107 23 20 2 3 55 6 7 66 1015 4 11 41 49 48 102 33 31 3 3 105 1 1 3 116 974 4 11 41 49 48 105 38 33 3 6 127 182 147 1238 4 11 41 49 48 107 34 31 2 3 55 7 8 72 1083 4 11 41 49 48 107 32 27 3 5 132 2 0 5 169 1213
4 11 41 49 48 107 30 27 4 5 80 11 96 1145
5 10 49 50 56 105 36 32 2 4 100 1 3 5 120 112
5 10 49 50 56 107 35 30 2 4 12 1 6 2 134 1206
5 10 49 50 56 107 27 29 1 4 86 11 1 105 10 57
5 10 49 50 56 107 32 29 2 2 74 8 9 85 1047
5 10 49 50 56 107 37 33 1 4 152 288 226 12 74
5 10 49 50 56 105 30 32 2 8 135 2 1 6 190 1136
6 12 49 50 56 107 25 20 2 3 24 3 8 29 1 3 1
6 12 49 50 56 104 30 28 5 2 95 136 130 1061
6 12 49 50 58 108 32 31 3 4 90 123 103 1194
6 12 49 50 58 108 30 29 2 4 110 159 143 1111
6 12 49 50 56 106 33 37 3 4 185 259 220 11 71
6 12 49 50 58 106 28 30 3 5 124 182 152 1197
7 10 50 54 57 106 39 34 3 7 114 1 7 3 151 11 45
7 10 50 54 57 109 25 31 2 6 102 146 125 1188
7 10 50 54 57 109 24 26 3 4 55 8 9 76 1171
7 10 50 54 57 109 36 33 2 5 113 158 128 12 18
7 10 50 54 57 109 37 33 3 7 100 1 9 8 105 1197
7 10 50 54 57 109 30 31 2 5 115 1 6 135 11 55
8 12 49 51 58 102 32 27 3 4 120 178 150 1186
8 12 49 51 58 108 32 35 3 5 85 11 9 106 11 22
8 12 49 51 58 102 22 18 2 1 19 3 21 1428
8 12 49 51 58 107 32 31 2 5 203 244 193 1264
8 12 49 51 58 107 40 39 6 7 124 185 137 135
9 11 42 47 48 101 34 30 2 5 101 148 115 12%
9 11 42 47 48 105 32 23 2 1 54 7 5 62 1209
9 11 42 47 48 105 33 29 3 8 145 193 165 11 69
A- lndlvldual plant data of F l gonontlon of the cmrr ICC 5718 x ICC 17101
En0 IPS DFF DSOXF DFP DM HT WD PB SB NPP Yld-P NSP HSW 2 9 5 45 49 5 3 5 107 3 6 5 2765 2 8 5 1 9 1 5 1 3 1 5 1 0 6 1 2125 3 1 0 5 4 5 5 50 53 1 0 7 5 3 6 7 3 0 2 3 7 4 9 7 7 1 3 6 133 1 2 4 5 4 9 5 47 49 5 6 5 109 35 3 1 2 3 4 2 116 1 3 5 1 3 1 5 1 0 0 3 5 9 5 4 5 5 49 5 7 5 108 3 2 8 30 2 8 3 8 8 9 8 1 1 7 6 9 6 1 1 2 2 6 8 9 5 4 8 5 505 5 8 5 104 34 24 2 5 4 84 1125 99 1 1 3 4 7 9 4 8 5 505 5 6 1 0 6 5 3 4 5 26 3 5 4 5 1 1 1 5 1 6 7 5 1 4 1 5 1 1 8 1 6 9 5 45 4 6 5 5 2 5 105 5 3 6 5 2 9 5 3 7 6 136 1 9 8 7 186 7 1 2 9 5 9 1 0 5 47 51 5 4 5 1 0 4 5 3 4 2 2 5 2 2 7 5 5 9 4 7 1 4 5 5 1 0 9 5 1 3 7 7
10 9 5 4 8 5 5 0 5 5 8 5 1025 37 34 4 5 6 9 8 2 1 3 3 5 1 0 7 2 1 2 4 9 11 11 42 50 49 1 0 4 5 3 4 5 2 6 2 2 5 4 7 99 1 5 5 7 1 3 2 2 1 1 8 6 12 8 5 46 5 0 5 57 1055 31 2 2 7 7 2 7 5 5 82 11 2 9 4 2 11 99 13 9 5 49 50 58 106 36 3 3 2 3 5 4 5 1 2 3 2 1 7 5 1 4 1 5 1 2 3 7 14 10 4 5 5 5 0 5 52 105 3 6 7 3075 3 4 2 8 2 5 1 9 6 5 1 5 6 2 11 35 15 3 5 55 6 0 5 6 7 5 109 3 1 5 30 3 4 8 3 5 1 1 7 5 9 2 5 1 2 4 1 16 8 45 5 0 5 6 2 5 107 3 9 7 2625 3 2 5 7 1 0 2 7 16 132 1 2 2 4 17 6 4 5 5 47 52 108 36 31 2 5 5 130 1585 1 4 0 5 11 56 18 7 51 5 4 5 5 9 5 1 1 0 5 3 9 2 3 4 2 2 7 5 2 1 0 9 7 1445 1 1 7 2 1 2 1 7 19 7 5 4 7 5 50 54 106 38 2 1 5 2 5 7 8 2 5 13 117 1 0 6 20 10 4 6 5 505 5 6 5 1035 38 3 5 1 2 5 5 5 1365 1 8 4 137 1 3 2 8 21 9 5 45 4 7 5 52 105 4 0 7 2 9 7 3 7 5 7 1 1 1 7 1 8 3 1 2 9 2 1 1 9 1 22 7 5 46 4 9 5 5 7 5 1 0 8 5 3 6 5 2 6 5 3 6 5 110 1 4 6 5 126 1 1 9 7 23 9 5 4 7 5 49 58 1 0 6 5 3 4 5 2 9 1 2 6 4 6 1 1 5 9 1 7 4 1 4 6 5 1 1 9 1 24 1 0 5 4 9 5 5 2 5 5 6 5 106 3 3 7 2 6 2 2 5 4 7 8 8 5 1 2 0 7 92 1345
25 9 5 42 48 4 9 5 1 0 6 5 31 19 3 5 5 81 1 1 5 101 1137 26 9 5 4 5 5 46 5 2 5 1 0 4 5 3 4 5 31 2 5 5 9 9 7 1 5 4 7 1 3 2 7 1 1 6 3 27 9 5 49 57 5 56 108 32 5 28 3 4 5 88 7 14 2 108 7 13 72
28 7 8 49 52 57 108 3 7 2 3 1 2 3 2 5 2 9 9 2 1 3 4 5 1 1 0 7 1 3 2
29 7 5 4 8 5 5 0 5 58 1045 3 5 5 32 3 2 4 5 1 4 1 5 1 6 1 7 1 5 1 8 1 1 9 7
30 10 51 52 5 6 5 108 3 5 2 30 3 5 6 2 1 1 3 7 17 1 3 8 5 1 1 4 6 31 7 5 43 46 5 2 5 1045 3 3 7 2 6 5 3 2 5 7 9 6 5 1525 1 2 5 5 1 2 4 1
32 11 42 4 7 5 4 8 5 1015 37 2 8 2 3 5 4 7 1015 1 5 3 7 1 2 4 5 1 2 9 1
33 9 5 47 51 57 1075 36 2 6 2 3 4 5 96 1 3 5 1 0 9 2 1 2 3 6
34 10 4 7 5 4 9 5 54 104 40 3 6 5 3 5 6 1155 1 7 7 144 1 2 2 4
35 8 4 9 5 54 58 105 2 8 2 22 3 5 6 91 2 1 2 0 2 9 9 2 5 1 2 0 1
36 9 5 45 49 53 107 5 34 5 24 7 3 5 4 7 92 5 12 87 120 12 1
37 1 0 5 44 49 5 3 5 106 3 4 2 28 2 7 6 2 9 1 7 1 3 1 2 110 1 1 9 2
38 9 5 4 8 5 51 5 7 5 1 0 5 5 3 6 5 2 3 7 3 7 4 5 9 1 7 5 1 1 7 5 9 3 7 1254
39 8 5 4 5 5 50 54 1 0 7 5 3 6 4 2 7 4 3 5 5 3 9 5 6 1 5 3 6 1 0 6 3 1 4 1
40 9 5 48 50 5 6 5 1 0 7 5 3 5 2 2 9 2 3 2 4 7 214 2072 1835 1 1 6 2
41 9 50 5 1 5 5 9 5 1 0 5 5 3 5 5 2 8 5 2 7 6 1 2 4 2 1 7 3 1 5 4 7 1 1 3 7
42 8 5 48 5 0 5 58 102 37 31 3 4 5 1 2 5 5 12 35 151 11 48
43 9 45 4 9 5 53 105 41 3 2 5 2 5 8 116 1 6 4 5 1325 1244
44 8 4 5 5 4 7 5 53 1045 3 4 2 2 9 2 2 7 5 5 91 7 1 5 8 2 108 1 4 8 9
45 7 48 51 5 6 5 1 0 6 5 3 2 5 27 3 5 8 8 2 1 3 2 5 1 0 2 5 1 2 8 3
48 8 5 52 53 60 110 43 3 0 2 3 2 6 5 1 3 3 2 2 1 4 2 1 4 9 2 1431
47 1 0 5 42 4 7 5 48 1 0 5 5 37 2 2 7 3 5 9 3 5 1 3 2 7 6 6 7 1 2 6 8
48 9 5 44 4 9 5 53 1 0 6 5 3 9 5 3 0 5 3 5 4 5 136 2 0 6 5 165 1 2 5 3
49 10 43 4 6 5 50 107 3 8 3 2 6 1 3 1 5 8 1 1 0 1 1 5 4 8 1 3 2 3 11 39
5" 9 5 46 4 9 5 54 103 3 6 5 3 1 5 3 2 6 5 1 2 9 3 1 8 6 7 1 6 2 5 1 1 3 6
51 1 0 5 4 8 5 5 0 5 57 107 3 5 2 6 2 5 2 7 6 5 103 1 4 0 2 1 1 0 7 1255
Indlvldual plan1 dab of dlllennl genofyps for w h b R o w r solour
En0 IPS OFF DsOX DFP DM HT WD PB SB NPP YldlP NSP HSW 1 8 5 47 5 9 5 49 1135 44 40 4 5 4 5 9 6 5 1005 4 7 1 2235 2 7 5 3 3 5 4 2 5 41 1075 34 3 5 5 3 4 5 8 3 5 9 5 2 1 1 5 2233 3 7 5 38 46 46 1 1 1 5 4 0 5 3 6 5 45 4 5 8 5 8 5 1 7 9 5 2002 4 8 5 42 5 0 5 4 9 5 111 32 295 4 5 4 5 69 9 5 5 1615 1679 5 1 0 5 30 49 3 5 5 111 33 31 3 2 5 465 4 4 5 9 5 2149 6 5 5 51 62 58 113 3 8 5 44 5 2 108 131 1685 1443 7 8 5 66 70 5 76 113 5 4 6 5 36 3 5 4 5 76 72 1085 1236 8 8 5 30 38 37 1 0 9 5 3 5 5 31 2 5 2 72 7 8 5 1 4 7 1924 9 8 5 33 4 3 5 3 9 5 115 41 33 2 5 5 5 1 0 5 1 1 4 5 2 0 7 1898 1 0 10 30 33 37 109 34 29 2 1 555 53 5 1075 21 31 11 10 38 47 47 113 33 31 5 3 5 3 755 82 1245 1431 1 2 9 5 2 7 5 245 5 3 5 1095 28 3 0 5 3 5 3 565 54 1 2 5 2316 1 3 5 4 9 5 51 5 7 ' 1 1 3 5 4 4 5 47 6 5 5 58 74 8 1093 1 4 6 2 7 5 5 0 5 33 111 4 3 5 38 4 4 126 1335 2295 3373 1 5 8 2 9 5 40 3 6 5 109 35 2 9 5 2 5 3 49 4 8 5 8 1 1947 1 6 6 36 5 2 5 43 112 5 42 34 4 2 5 77 5 7 6 5 12 55 1648 1 7 8 5 36 48 4 2 5 114 33 265 3 3 58 70 1 0 2 1492 1 8 9 M 5 7 2 5 73 1135 49 42 35 7 5 1 0 5 5 1 0 4 1 5 9 2 1538 1 9 6 48 51 56 110 40 3 6 5 4 3 5 88 96 16 1505 20 9 48 51 56 110 40 4 0 5 4 3 5 68 96 16 1505 21 1 0 4 8 5 6 0 5 5 5 5 1085 40 3 3 5 5 4 5 1 0 1 5 1 0 6 5 1 7 7 5 1885 22 10 27 3 2 5 33 1 1 0 2 3 5 32 2 5 1 5 3 7 5 405 7 2 5 1 8 0 1 23 6 50 57 57 5 1135 4 9 5 44 3 5 8 68 705 1205 17 2 24 9 5 5 6 5 8 6 5 64 111 3 5 5 3 5 5 3 4 5 85 1005 1 8 3 1673 25 8 4 3 5 4 9 5 5 2 5 110 40 3 4 5 3 3 5 9 2 5 1005 1635 1622 26 1 0 6 0 5 70 6 9 5 111 42 3 5 5 2 5 4 5 7 6 5 7 5 1 2 0 5 1655 27 7 34 42 3 7 5 112 42 36 3 3 78 7 8 5 1 4 1 1821 28 8 5 4 6 5 6 6 5 53 1 1 1 5 4 8 5 46 4 7 1 0 2 5 113 17 1519 29 9 6 0 5 50 38 1105 45 3 9 5 3 6 2 0 6 1 2 0 3 1 1 5 2 0 9 30 6 30 4 4 5 3 6 5 1 1 0 4 4 5 38 3 2 5 8 2 5 7 6 5 1 5 4 2 2 0 5 2 31 8 5 26 34 33 1105 40 40 5 5 5 1 3 7 5 1 1 8 1 6 2 5 1035 3 2 7 3 5 5 82 41 5 113 5 0 5 48 4 5 119 110 21 05 1975 33 6 3 6 5 56 4 3 5 1105 45 39 3 5 5 137 138 2 0 7 15 34 7 5 40 5 6 5 47 113 39 38 3 2 5 665 6 3 5 1 2 4 5 1 8 9 35 7 38 465 45 111 295 31 5 3 2 5 65 183 1 6 5 1909 36 11 45 5 5 5 5 2 5 1125 4 0 5 39 2 3 81 8 1 5 1 6 5 2003 37 9 5 31 5 0 5 3 7 5 1115 35 3 4 5 3 2 5 7 2 5 6 8 5 1 5 2 2 2 9 38 8 5 32 42 3 9 5 109 36 33 2 5 3 5 1155 1365 2 1 1 1561 39 11 3 4 5 4 8 5 4 0 5 111 3 5 5 3 4 5 4 4 8 7 5 6 2 5 1043 2277 4 0 10 37 5 45 45 1095 3 0 5 3 0 5 3 3 5 1085 64 5 13 1 2046 41 8 5 36 45 43 1125 38 3 3 5 3 2 91 1 0 0 1 5 9 5 1573 42 9 5 41 5 51 4 8 5 111 5 43 455 4 4 5 215 118 1 8 3 1529 4 3 8 5 31 50 3 7 5 111 46 45 3 5 2 5 86 85 1855 1 9 8 4 4 6 32 38 3 7 5 1075 3 0 3 2 5 2 5 2 5 7 1 5 9 5 1 5 2 5 1732 4 6 9 43 85 5 1 5 1 1 1 5 3 1 5 3 1 5 4 5 3 5 91 8 6 5 2 0 1 5 2 3 3 3 4 7 4 5 3 1 5 U 37 1 1 0 5 3 0 5 30 3 4 98 8 7 5 1 2 4 5 1072 48 6 5 29 47 30 1085 3 0 5 38 5 2 4 62 5 60 1 3 2 22M1 49 7 2 6 5 34 3 3 5 113 5 0 4 8 5 4 4 5 131 9 1 1 2 8 5 137 50 9 48 5 1 5 5 5 5 112 40 34 4 3 5 6 1 5 7 4 5 9 8 1 4 7 8 51 8 29 34 3 5 1 0 8 5 27 26 2 1 5 45 4 0 8 6 1 1958 52 9 52 8 3 5 5 9 5 111 4 6 4 2 5 4 7 5 197 1 8 8 3 3 8 5 1907
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