Theor Appl Genet (1984) 68:487-491
9 Springer-Verlag 1984
Line • tester analysis for fixed effect model in cotton ( Gossypium hirsutum L.) L. S. Kaushik, D. P. Singh and R. S. Paroda Department of Plant Breeding, Haryana Agricultural University, Hisar-125 004, India Received January 10, 1983; Accepted February 20, 1984 Communicated by A. Robertson
Summary. The data from an experiment in cotton consisting of three testers and 12 lines selected deliberately have been analysed. The investigation showed higher specific combining ability variance for yield o f seed cotton and n u m b e r of bolls, indicating the predominance of non-additive gene action. O f parental lines, H777 was found to possess high g.c.a, effects for seed cotton yield, n u m b e r o f bolls and n u m b e r of sympodes. Parent H842 contributed only for boll weight, whereas H655 was good general combiner for n u m b e r o f monopodes. There appeared to be better chances for increasing the yield by exploiting hybrid vigour for the n u m b e r o f bolls and boll weight. The presence o f marked non-additive gene effects, in addition to additive gene effects, indicated the need for exploiting both the fixable and non-fixable components o f genetic variance for increasing productivity in cotton.
Key words: Combining ability analysis - Gossypium
hirsutum L.
Introduction Combining ability analysis is the most widely used biometrical tool for identifying prospective parents and for formulating breeding procedures most likely to succeed. Line x tester analysis, which is a simple extension and application o f the analysis o f two factor factorial experiments given by Fisher (1926) and Yates (1937), has been very c o m m o n l y used for combining ability analysis in plant breeding. With the above objective in mind, a line X tester design was adopted in cotton (Gosaypium hirsutum L.) and the analysis presented with the help o f experimental data.
Material and methods Three testers H777, H655 and H842 and 12 lines (females) BARSP84, REBA-B50, BP68, Laxmi 7882, PL807, H807, 152F, Paymaster-303, Cocker310, SV63, G.S.29 and H843 were deliberately selected, keeping in view the phenotypic and geographic diversity. Fifteen parents and 36 crosses were grown in a Randomized Block Design (RBD) with three replications during kharif 1981 at the Haryana Agricultural University Farm, Hisar. Row to row spacing was kept at 75 cm and plant to plant spacing was 30 cm. Ten competitive plants were selected from Fx's and parents. Data was recorded on yield per plant (gm), boll number per plant, boll weight (gm), the number of monopodes and sympodes. Regarding statistical analysis, the combined analysis of parents and crosses was done as suggested by Arunachanlam (1974) and for combining ability analysis, the following model was used: Yijk= Ix+ fi+ mj + (mOij + bk + eijk (i= 1, 2. . . . 1,j = 1, 2 . . . . t, k = 1, 2 . . . . r) where Yijk denotes the observation recorded on the (i• cross in the k th replication; ~t is the general effect; fi is the effect of the i th line; mj is the effect of the j th tester; sij is the specific combining ability (s.c.a.) effect of the (i xj)th cross; bk is the k th block effect and eijk is the environmental effect (random error) associated with the (ijk)th observation which is assumed to be normally and independently distributed with a mean of zero and variance (o2). The estimates of male, female and interaction effects are obtained by using the usual least square theory. Male and female effects are obtained by taking the deviation of respective means from the overall mean, and the interaction effect is obtained after subtracting male and female effects from the cross effect. The sum of squares due to effects are obtained by the method of fitting constants (Table 1). 6~, 6~n represent fixed type variances for lines and testers respectively. The weighted average of these two can be taken as an estimate of #~.c.a., i.e. #~.c.a.= [(1 - 1) cr}+ (t - 1) a~n]/(1 + t - 2). The ratio t~Z~.c.a./fs2.c.a,will give an approximate idea about degree of dominance.
488
L.S. Kaushik et al.: Line x tester analysis for fixed effect m o d e l in cotton
Table 1. A N O V A table for line x tester analysis (fixed effect m o d e l ) Source
d.f.
Sum o f squares
Blocks
r- 1
Lines
1- 1
Y2.k
y2
It
ltr
y2..
y2 ... ltr
K
i E y2j.
t- l
Line x tester
(1 - 1) x (t -- 1)
J
2y2j. "J r y2.
Mj
0-2 + r t Y', f2/(1 - 1) i
Mr/Me
ltr
Mt
o"2 + rl ~ m ~ / ( t - 1) j
Mt/Me
~'~ Y~.. i rt
MIx t
0"2 q"
Me
0-2
i 2(mf){j "J (1 -- 1) (t -- 1)
MI • t/Me
y2 q - - _ _ .,.
lr (It - 1 ) x ( r - 1)
F
y2 ""
rl
j
Error
Expexted m e a n square
...
rt Testers
M.S.
~ Y~jk i,j,k
ltr -- ~ Y .2k m K
- ~, y2j./r + Y 2 . / l t r LJ
where Yi... = ~, Yijk, Y.j. = ZYijk, Y...= ~, Yijk and Y..k = Z Yijk j,k
i,k
i,j,k
~,j
To evaluate the relative contribution o f g.c.a, and s.c.a., fixed type o f variances are obtained as follows: if} = ~, f2/(1 - 1) = (M 1- Me)/r t i
if2m = ~'~ mY/(t - 1) = (M t - M e ) / r l J ~s2c-a = a2mxf= 2 (m f){J/(1 - 1) (t - 1) = (MI• t - M e ) / r ~-2 = Me ~'J Table 2. A N O V A table for parents and crosses Source
d.f.
Replications Hybrids Parents Hybrids vs parents Error
2 35 14 1 100
Total
152
Mean square Yield
No. o f bolls
Boll wt
No. o f m o n o p o d e s
No. o f s y m p o d i a
2.338 1280.037"* 769.209** 2790.275** 57.607
99.261 468.207** 297.305** 1161.956"* 17.371
0.1182 0.976** 1.266"* 0.683** 0.055
0.082 1.343"* 4.835** 1.563"* 0.208
2.789 37.526** 40.649** 1.418NS 2.619
Yield
No. o f bolls
Boll wt
No. o f m o n o p o d e s
No. o f s y m p o d e s
112.5156 1615.8781 ** 1031.7215"* 1134.6920** 54.3963**
146.0417 731.3633"* 77.4559** 372.1526"* 15.2492
0.2420 2.5337** 0.3306** 0.2563** 0.0401
0.0395 3.3293** 1.2993"* 0.3540 0.2310
0.7612 61.8561"* 2.9515"* 28.5044** 2.3994
Table 3. A N O V A for combining ability analysis Source
Replication Line (female) Tester (males) Line x tester Error Total
d.f.
2 11 2 22 70 107
Mean squares
L. S. Kaushik et al.: Line x tester analysis for fixed effect model in cotton
Results and discussion The analysis o f v a r i a n c e (Table 2) o f the p a r e n t s a n d crosses for five characters, viz. yield o f seed cotton, n u m b e r o f bolls, boll weight, n u m b e r o f m o n o p o d i a a n d n u m b e r o f s y m p o d i a i n d i c a t e d that n o significant differences existed b e t w e e n replications, w h e r e a s differences b e t w e e n hybrids, p a r e n t s a n d h y b r i d s vs p a r e n t s w e r e significant for all c h a r a c t e r s e x c e p t n u m ber of sympodes. A n a l y s i s o f v a r i a n c e for c o m b i n i n g ability ( T a b l e 3) r e v e a l e d that the differences d u e to f e m a l e a n d m a l e p a r e n t s w e r e h i g h l y significant for all c h a r a c t e r s s t u d i e d e x c e p t d u e to m a l e s for the n u m b e r o f s y m p o d e s . C o n tribution of male• i n t e r a c t i o n was h i g h l y significant for all c h a r a c t e r s e x c e p t n u m b e r o f m o n o p o d e s . The differences d u e to m a l e s a n d females, w h e n tested against interaction, as is d o n e in a r a n d o m effect m o d e l , turns out to be n o n - s i g n i f i c a n t w h i c h is c o n t r a r y to the c o n c l u s i o n d r a w n f r o m a fixed effect m o d e l . H e n c e , g e n e effects t h o u g h p r e s e n t are u n d e r - e s t i m a t e d or n o t
489
d e t e c t e d in a r a n d o m effect analysis w h e n actual d a t a pertains to a fixed effect m o d e l . The estimates o f O-g.c.a .-2 and ~rsca. ~~ g i v e n in T a b l e 4 r e v e a l e d that the n a t u r e o f the g e n e effects was p r e d o m i n a n t l y n o n - a d d i t i v e for yield, n u m b e r o f bolls a n d sympodes. However, substantial additive gene action was n o t e d for boll w e i g h t a n d the n u m b e r o f m o n o podes. The c o m p a r a t i v e analysis o f gca effects ( T a b l e 5) o f the p a r e n t s s h o w e d that a m o n g tester p a r e n t s o n l y H777 was a b e t t e r g e n e r a l c o m b i n e r for seed c o t t o n yield a n d the n u m b e r o f bolls. O n l y f o u r females, B A R - S P - 8 4 , L a x m i 7 8 8 2 , PL807 a n d H807, s h o w e d c o n s i d e r a b l e positive g.c.a, effects for three i m p o r t a n t traits-yield, boll n u m b e r a n d s y m p o d i a - g i v i n g little choice a m o n g f e m a l e s with respect to g.c.a, effects. W i t h r e g a r d to s.c.a, effects (Table 6) 17 cross c o m b i n a t i o n s s h o w e d positive estimates o f s.c.a, effects for the yield o f seed cotton. O f these h y b r i d s H842 x 152F, H655 x H807, H777 x H807, H842 x P a y m a s t e r 303 a n d H842 X SV63, in that order, w e r e the best five c o m b i n a -
Table 4. Estimation of fixed type variances for different traits Estimate of variances (fixed type)
Yield
Boll wt
Boll no.
No. of monopodes
No. of sympodes
62 = M e #2c.a. = (M1 x t - Me)/3 6~ = ( M ~ - Me)/9 62 = (Nit- Me)/36 62.c.a = (11 62+2 62)/13 62c.a/62.c.a"
54.3963 360.0986 173.4980 27.1479 150.9826 2.3850
0.0401 0.0721 0.2771 0.0081 0.2357 0.3059
15.2492 118.9678 79.5682 1.7280 67.5928 1.7601
0.2310 0.0410 0.3443 0.0297 0.2959 0.1386
2.3994 8.7017 6.6063 0.0153 5.5923 1.5560
Table 5. g.c.a, effects of parents for different characters Sr. no.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Parent Males H777 H655 H842 Females BAR-SP-84 REBA-B-50 BP68 L7882 PL807 H807 152F Paymaster 303 Cocker 310 S.V.63 G.S.29 H843
Yield
Boll no.
Boll wt
No. of monopodes
No. of sympodes
6.17 - 2.76 - 3.41
1.585 - 0.276 - 1.309
0.038 - 0.109 0.070
0.126 0.092 - 0.219
0.293 - 0.013 - 0.279
8.05 - 19.75 - 13.43 9.65 26.25 14.97 - 5.85
6.485 12.715 - 2.804 8.752 13.330 12.096 1.763
- 0.400 0.605 - 0.596 - 0.249 - 0.049 - 0.306 - 0.517
-
0.798 0.169 0.191 0.413 0.313 0.124 0.591
2.096 - 2.715 - 1.926 4.429 3.918 3.057 - 0.349
5.73 - 6.97 - 13.45 0.85 - 6.02
- 7.004 - 10.693 - 5.315 4.352 - 8.248
0.842 0.816 - 0.249 - 0.366 0.467
- 0.258 - 0.102 - 0.313 1.598 0.076
-
2.149 1.738 2.349 1.060 1.204
490
L.S. Kaushik et al.: Line • tester analysis for fixed effect model in cotton
Table 6. s.c.a, effects of different crosses for different characters Cross
Yield
Boll no.
Boll wt
No. of monopodes
No. o f s y m p o d e s
1• 4 1• 5 1• 7 1x 8 1• 9
12.27 - 11.10 - 11.82 - 0.70 10.44 23.65
1 • 10
-29.00
1• 1x 1• 1• 1• 2• 2• 2• 2• 2• 2• 2• 2x 2x 2• 2• 2• 3x 3• 3• 3x 3• 3• 3• 3• 3• 3• 3• 3•
-6.71 13.02 - 7.63 5.80 1.74 4.33 - 2.47 2.53 1.50 - 3.77 26.48 -2.00 - 15.28 - 2.45 - 6.90 4.20 -6.26 - 16.62 13.55 9.26 - 0.82 -6.68 - 50.14 30.98 22.00 - 10.56 14.51 - 10.02 4.52
3.493 - 2.707 - 6.185 - 4.507 10.281 15.915 - 14.418 - 7.585 - 0.062 - 2.040 7.226 0.593 4.521 - 1.646 0.676 0.954 5.109 11.310 - 2.424 - 3.657 1.565 - 9.146 - 3.980 - 3.280 - 8.013 4.384 5.509 3.553 - 15.391 - 27.224 16.842 11.243 - 1.502 11.187 - 3.247 2.687
-0.067 0.131 -0.455 0.326 -0.172 -0.128 -0.194 0.254 0.483 - 0.212 -0.191 0.202 0.140 0.028 0.126 -0.038 -0.281 0.279 0.030 -0.182 - 0.356 0.152 0.289 -0.187 -0.073 -0.158 0.330 - 0.314 0.453 -0.150 0.164 - 0.071 -0.125 0.060 - 0.097 - 0.013
- 0.104 0.163 - 0.415 - 0.326 -0.126 0.185 0.418 0.519 0.029 - 0.260 -0.071 - 0.015 0.263 -0.570 -0.098 0.108 0.308 -0.219 -0.181 -0.381 0.163 0.274 - 0.203 0.052 -0.073 0.408 0.463 0.219 -0.181 -0.404 - 0.237 - 0.136 -0.192 -0.015 0.247 - 0.037
3.796 0.207 - 0.949 - 1.071 2.440 3.074 - 4.193 -0.259 - 1.504 0.441 - 0.982 - 1.004 1.102 - 1.421 - 1.510 2.069 0.113 3.147 - 2.120 - 0.287 1.002 0.113 - 0.942 - 1.265 - 0.159 1.212 2.456 - 0.999 - 2.554 -6.221 6.313 0.546 0.502 - 0.554 1.924 2.268
1•
6
11 12 13 14 15 4 5 6 7 8 9 10 11 12 13 14 15 4 5 6 7 8 9 10 11 12 13 14 15
tions. It is i n t e r e s t i n g to n o t e t h a t all five crosses s h o w i n g h i g h e r s.c.a, effects for y i e l d o f s e e d c o t t o n also r e c o r d e d t h e g r e a t e r v a l u e s o f s.c.a, effects for b o l l n u m b e r . T h e s.c.a, for y i e l d o f s e e d c o t t o n a n d boll n u m b e r h a d a l m o s t a p a r a l l e l t r e n d . S i x t e e n cross c o m b i n a t i o n s e x h i b i t e d p o s i t i v e e s t i m a t e s o f s . c . a , effects for boll w e i g h t , t h e m a x i m u m b e i n g in case o f H 7 7 7 x C o c k e r 310, (0.483) f o l l o w e d b y H 8 4 2 x P L 8 0 7 (0.453), H842 • BP68 (0.33), H777 • L a x m i 7882 (0.326). E i g h t e e n c r o s s e s p o s s e s s e d p o s i t i v e s.c.a, effects for n u m b e r o f s y m p o d e s , H 8 4 2 x 152F h a v i n g the m a x i m u m s.c.a. v a l u e (6.313) c l o s e l y f o l l o w e d b y H777 x B A R SP84 (3.796), H655 x H807 (3.14) a n d H777 • H807 (3.074). H y b r i d H842 x 152F, p o s s e s s i n g t h e h i g h e s t s.c.a, effect for n u m b e r o f s y m p o d i a , also p o s s e s s e d t h e h i g h e s t s.c.a, effect for s e e d c o t t o n yield.
Heterotic effects H e t e r o t i c effects w e r e e s t i m a t e d as the d e v i a t i o n s o f the Fl's from m i d - p a r e n t a l values (MP) and the deviations
f r o m its b e t t e r p a r e n t (BP), w i t h r e s p e c t to five traits o f A m e r i c a n cotton. S i x t e e n crosses m a n i f e s t e d h e t e r o s i s o v e r t h e m i d p a r e n t for s e e d c o t t o n yield. C r o s s H842 x H807 (69.81) f o l l o w e d b y H 7 7 7 X P L 8 0 7 (61.81) a n d H 8 4 2 • (61.57) e x h i b i t e d e v e n m o r e t h a n 50% h e t e r o s i s o v e r the mid-parent. Only eight hybrids showed positive h e t e r o s i s o v e r the b e t t e r p a r e n t for s e e d c o t t o n yield, T w o cross c o m b i n a t i o n s , H 7 7 7 • (42.9%) a n d H777 X PL807 (40.2), r e c o r d e d m o r e t h a n 40% h e t e r o s i s o v e r t h e b e t t e r p a r e n t (H777), t h e m o s t c o m m o n l y c u l t i v a t e d c o t t o n v a r i e t y o f the z o n e . Similarly, 16 crosses e x h i b i t e d p o s i t i v e h y b r i d v i g o u r ( M P ) for n u m b e r o f bolls, r a n g i n g b e t w e e n 8.0 to 85.2 (H655 x H807) p e r cent. T w e l v e crosses r e c o r d e d h e t e r o s i s o v e r BP for boll n u m b e r . H y b r i d s H655 x H807 (74.0%), H 6 5 5 • (53.5%) a n d H 6 5 5 • B A R SP (51.4%), r e c o r d e d m o r e t h a n 50% h e t e r o s i s o v e r BP for boll n u m b e r . E l e v e n crosses w e r e f o u n d to b e s u p e r i o r for boll w e i g h t o v e r t h e i r b e t t e r p a r e n t s .
L. S. Kaushik et al.: Line • tester analysis for fixed effect model in cotton Heaviest boll weight was p r o d u c e d b y the crosses H777 x Laxmi 7882 (16.5%), followed by H842 X PL807 (11.9%) and H 7 7 7 x H 8 0 7 (10.5%). More than 50% h y b r i d vigour (BP) was recorded for n u m b e r o f m o n o podes in two crosses, H842 x BP68 (52.2%) and H842 x Reba-B50 (50.8%). Only six crosses recorded h i g h e r n u m b e r o f s y m p o d i a (BP), ranging between 3.0% (H655 X H807) to 28.0% (H777 X H807). The degree of heterosis varied considerably for different characters with the maximum heterosis for the number of bolls per plant, yield of seed cotton and boll weight. Therefore, there appears to be good possibilities of increasing the yield by exploiting hybrid vigour through the number of bolls and boll weight. These results are in confirmity with earlier reports (White and Richmond 1963; Miller and Marani 1963; Marani 1967; Singh and Murty 1971; Singh 1974). For yield of seed cotton, the three best crosses were H842 x 152F, H655XH807 and H777 x H807. These crosses were also best for boll number and the number of both types of branches. Crosses involving parents H842, H807 and 152F exhibited higher degree of heterosis for yield of seed cotton and other component characters of yield in this study. The best combinations (H777xPL807 and H777XH807), as judged from the s.c.a. estimates of seed cotton, involved the best general combiners (Table 6). Therefore, there may be a good chance of success for getting high yielding segregants and it equally may be possible of developing a hybrid involving the parents H777, PL807 and H807. These results are in general agreement with those reported by Turner (1953); Marani (1963, 1964); Gururaja Rao (1975) and Bhallala (1976). In order to exploit the type of gene effects operative in the population developed by involving H777, H807 and PL807, a breeding procedure which takes care of both additive and non-additive gene effects might prove the most efficient in improving the population. Similar findings have also been reported by Gupta and Singh 1970; Singh 1974; Chandermathi and Menon 1973 and Mirza and Khan 1976.
491
References Arunachalam VC (1974) The fallacy behind the use of a modified line x tester design. Indian J Genet 34:280-287 Bhallala MK (1976) Heterosis analysis in upland cotton. MSc Thesis (submitted to HAU, Hisar) Chanderamathi PS, Menon PM (1973) Diallel analysis of yield and its components in G. barbedensi L. Madras Agric J 60: 1560-1566 Fisher RA (1926) The arrangement of field experiments. J Min Agric 33:503-513 Gupta MP, Singh RB (1970) Genetics of some characters in upland cotton. Indian J Genet 30:590-598 Gururajarao MR (1975) Genetic analysis in upland cotton (G. hirstu L.). Mysore J Agric Sci 9:199 Marani A (1963) Heterosis and combining ability for yield and its components in cotton. Crop Sci 3:552-555 Marani A (1964) Heterosis and combining ability in diallel cross of two spps. of cotton. Crop Sci 4:265-268 Marani A (1967) Heterosis and combining ability in intra- and interspecific crosses of cotton. Crop Sci 7:519-522 Miller PA, Marani A (1963) Heterosis and combining ability in varietal top crosses of upland cotton (G. hirsutum L.). Crop Sci 3: 646-649 Mirza AM, Khan MA (1976) Diallel analysis of some important characters in cotton (G. hirsutum L.) Plant Breed (Abstr) 46:3509 Singh BB, Murty BR (1971) Hybrid vigour in intervarietal crosses of upland cotton. Indian J Genet 31:1-7 Singh DP (1974) Genetic architecture of Gossypium hirsutum for certain economic character. PhD Thesis (submitted to Int Adv Study Meerut) Turner JH (1953) A study of heterosis in upland cotton yield of hybrids compared with varieties. Agron J 45:484-486 Yates F (1937) The design and analysis of factorial experiments. Imp Bureau Soil Sci Tech Comm 35:1-95 White TG, Richmond TR (1963) Heterosis and combining ability in top and diallel crosses in cotton. Crop Sci 6:58-63
9 Springer-Verlag 1984
Line • tester analysis for fixed effect model in cotton ( Gossypium hirsutum L.) L. S. Kaushik, D. P. Singh and R. S. Paroda Department of Plant Breeding, Haryana Agricultural University, Hisar-125 004, India Received January 10, 1983; Accepted February 20, 1984 Communicated by A. Robertson
Summary. The data from an experiment in cotton consisting of three testers and 12 lines selected deliberately have been analysed. The investigation showed higher specific combining ability variance for yield o f seed cotton and n u m b e r of bolls, indicating the predominance of non-additive gene action. O f parental lines, H777 was found to possess high g.c.a, effects for seed cotton yield, n u m b e r o f bolls and n u m b e r of sympodes. Parent H842 contributed only for boll weight, whereas H655 was good general combiner for n u m b e r o f monopodes. There appeared to be better chances for increasing the yield by exploiting hybrid vigour for the n u m b e r o f bolls and boll weight. The presence o f marked non-additive gene effects, in addition to additive gene effects, indicated the need for exploiting both the fixable and non-fixable components o f genetic variance for increasing productivity in cotton.
Key words: Combining ability analysis - Gossypium
hirsutum L.
Introduction Combining ability analysis is the most widely used biometrical tool for identifying prospective parents and for formulating breeding procedures most likely to succeed. Line x tester analysis, which is a simple extension and application o f the analysis o f two factor factorial experiments given by Fisher (1926) and Yates (1937), has been very c o m m o n l y used for combining ability analysis in plant breeding. With the above objective in mind, a line X tester design was adopted in cotton (Gosaypium hirsutum L.) and the analysis presented with the help o f experimental data.
Material and methods Three testers H777, H655 and H842 and 12 lines (females) BARSP84, REBA-B50, BP68, Laxmi 7882, PL807, H807, 152F, Paymaster-303, Cocker310, SV63, G.S.29 and H843 were deliberately selected, keeping in view the phenotypic and geographic diversity. Fifteen parents and 36 crosses were grown in a Randomized Block Design (RBD) with three replications during kharif 1981 at the Haryana Agricultural University Farm, Hisar. Row to row spacing was kept at 75 cm and plant to plant spacing was 30 cm. Ten competitive plants were selected from Fx's and parents. Data was recorded on yield per plant (gm), boll number per plant, boll weight (gm), the number of monopodes and sympodes. Regarding statistical analysis, the combined analysis of parents and crosses was done as suggested by Arunachanlam (1974) and for combining ability analysis, the following model was used: Yijk= Ix+ fi+ mj + (mOij + bk + eijk (i= 1, 2. . . . 1,j = 1, 2 . . . . t, k = 1, 2 . . . . r) where Yijk denotes the observation recorded on the (i• cross in the k th replication; ~t is the general effect; fi is the effect of the i th line; mj is the effect of the j th tester; sij is the specific combining ability (s.c.a.) effect of the (i xj)th cross; bk is the k th block effect and eijk is the environmental effect (random error) associated with the (ijk)th observation which is assumed to be normally and independently distributed with a mean of zero and variance (o2). The estimates of male, female and interaction effects are obtained by using the usual least square theory. Male and female effects are obtained by taking the deviation of respective means from the overall mean, and the interaction effect is obtained after subtracting male and female effects from the cross effect. The sum of squares due to effects are obtained by the method of fitting constants (Table 1). 6~, 6~n represent fixed type variances for lines and testers respectively. The weighted average of these two can be taken as an estimate of #~.c.a., i.e. #~.c.a.= [(1 - 1) cr}+ (t - 1) a~n]/(1 + t - 2). The ratio t~Z~.c.a./fs2.c.a,will give an approximate idea about degree of dominance.
488
L.S. Kaushik et al.: Line x tester analysis for fixed effect m o d e l in cotton
Table 1. A N O V A table for line x tester analysis (fixed effect m o d e l ) Source
d.f.
Sum o f squares
Blocks
r- 1
Lines
1- 1
Y2.k
y2
It
ltr
y2..
y2 ... ltr
K
i E y2j.
t- l
Line x tester
(1 - 1) x (t -- 1)
J
2y2j. "J r y2.
Mj
0-2 + r t Y', f2/(1 - 1) i
Mr/Me
ltr
Mt
o"2 + rl ~ m ~ / ( t - 1) j
Mt/Me
~'~ Y~.. i rt
MIx t
0"2 q"
Me
0-2
i 2(mf){j "J (1 -- 1) (t -- 1)
MI • t/Me
y2 q - - _ _ .,.
lr (It - 1 ) x ( r - 1)
F
y2 ""
rl
j
Error
Expexted m e a n square
...
rt Testers
M.S.
~ Y~jk i,j,k
ltr -- ~ Y .2k m K
- ~, y2j./r + Y 2 . / l t r LJ
where Yi... = ~, Yijk, Y.j. = ZYijk, Y...= ~, Yijk and Y..k = Z Yijk j,k
i,k
i,j,k
~,j
To evaluate the relative contribution o f g.c.a, and s.c.a., fixed type o f variances are obtained as follows: if} = ~, f2/(1 - 1) = (M 1- Me)/r t i
if2m = ~'~ mY/(t - 1) = (M t - M e ) / r l J ~s2c-a = a2mxf= 2 (m f){J/(1 - 1) (t - 1) = (MI• t - M e ) / r ~-2 = Me ~'J Table 2. A N O V A table for parents and crosses Source
d.f.
Replications Hybrids Parents Hybrids vs parents Error
2 35 14 1 100
Total
152
Mean square Yield
No. o f bolls
Boll wt
No. o f m o n o p o d e s
No. o f s y m p o d i a
2.338 1280.037"* 769.209** 2790.275** 57.607
99.261 468.207** 297.305** 1161.956"* 17.371
0.1182 0.976** 1.266"* 0.683** 0.055
0.082 1.343"* 4.835** 1.563"* 0.208
2.789 37.526** 40.649** 1.418NS 2.619
Yield
No. o f bolls
Boll wt
No. o f m o n o p o d e s
No. o f s y m p o d e s
112.5156 1615.8781 ** 1031.7215"* 1134.6920** 54.3963**
146.0417 731.3633"* 77.4559** 372.1526"* 15.2492
0.2420 2.5337** 0.3306** 0.2563** 0.0401
0.0395 3.3293** 1.2993"* 0.3540 0.2310
0.7612 61.8561"* 2.9515"* 28.5044** 2.3994
Table 3. A N O V A for combining ability analysis Source
Replication Line (female) Tester (males) Line x tester Error Total
d.f.
2 11 2 22 70 107
Mean squares
L. S. Kaushik et al.: Line x tester analysis for fixed effect model in cotton
Results and discussion The analysis o f v a r i a n c e (Table 2) o f the p a r e n t s a n d crosses for five characters, viz. yield o f seed cotton, n u m b e r o f bolls, boll weight, n u m b e r o f m o n o p o d i a a n d n u m b e r o f s y m p o d i a i n d i c a t e d that n o significant differences existed b e t w e e n replications, w h e r e a s differences b e t w e e n hybrids, p a r e n t s a n d h y b r i d s vs p a r e n t s w e r e significant for all c h a r a c t e r s e x c e p t n u m ber of sympodes. A n a l y s i s o f v a r i a n c e for c o m b i n i n g ability ( T a b l e 3) r e v e a l e d that the differences d u e to f e m a l e a n d m a l e p a r e n t s w e r e h i g h l y significant for all c h a r a c t e r s s t u d i e d e x c e p t d u e to m a l e s for the n u m b e r o f s y m p o d e s . C o n tribution of male• i n t e r a c t i o n was h i g h l y significant for all c h a r a c t e r s e x c e p t n u m b e r o f m o n o p o d e s . The differences d u e to m a l e s a n d females, w h e n tested against interaction, as is d o n e in a r a n d o m effect m o d e l , turns out to be n o n - s i g n i f i c a n t w h i c h is c o n t r a r y to the c o n c l u s i o n d r a w n f r o m a fixed effect m o d e l . H e n c e , g e n e effects t h o u g h p r e s e n t are u n d e r - e s t i m a t e d or n o t
489
d e t e c t e d in a r a n d o m effect analysis w h e n actual d a t a pertains to a fixed effect m o d e l . The estimates o f O-g.c.a .-2 and ~rsca. ~~ g i v e n in T a b l e 4 r e v e a l e d that the n a t u r e o f the g e n e effects was p r e d o m i n a n t l y n o n - a d d i t i v e for yield, n u m b e r o f bolls a n d sympodes. However, substantial additive gene action was n o t e d for boll w e i g h t a n d the n u m b e r o f m o n o podes. The c o m p a r a t i v e analysis o f gca effects ( T a b l e 5) o f the p a r e n t s s h o w e d that a m o n g tester p a r e n t s o n l y H777 was a b e t t e r g e n e r a l c o m b i n e r for seed c o t t o n yield a n d the n u m b e r o f bolls. O n l y f o u r females, B A R - S P - 8 4 , L a x m i 7 8 8 2 , PL807 a n d H807, s h o w e d c o n s i d e r a b l e positive g.c.a, effects for three i m p o r t a n t traits-yield, boll n u m b e r a n d s y m p o d i a - g i v i n g little choice a m o n g f e m a l e s with respect to g.c.a, effects. W i t h r e g a r d to s.c.a, effects (Table 6) 17 cross c o m b i n a t i o n s s h o w e d positive estimates o f s.c.a, effects for the yield o f seed cotton. O f these h y b r i d s H842 x 152F, H655 x H807, H777 x H807, H842 x P a y m a s t e r 303 a n d H842 X SV63, in that order, w e r e the best five c o m b i n a -
Table 4. Estimation of fixed type variances for different traits Estimate of variances (fixed type)
Yield
Boll wt
Boll no.
No. of monopodes
No. of sympodes
62 = M e #2c.a. = (M1 x t - Me)/3 6~ = ( M ~ - Me)/9 62 = (Nit- Me)/36 62.c.a = (11 62+2 62)/13 62c.a/62.c.a"
54.3963 360.0986 173.4980 27.1479 150.9826 2.3850
0.0401 0.0721 0.2771 0.0081 0.2357 0.3059
15.2492 118.9678 79.5682 1.7280 67.5928 1.7601
0.2310 0.0410 0.3443 0.0297 0.2959 0.1386
2.3994 8.7017 6.6063 0.0153 5.5923 1.5560
Table 5. g.c.a, effects of parents for different characters Sr. no.
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Parent Males H777 H655 H842 Females BAR-SP-84 REBA-B-50 BP68 L7882 PL807 H807 152F Paymaster 303 Cocker 310 S.V.63 G.S.29 H843
Yield
Boll no.
Boll wt
No. of monopodes
No. of sympodes
6.17 - 2.76 - 3.41
1.585 - 0.276 - 1.309
0.038 - 0.109 0.070
0.126 0.092 - 0.219
0.293 - 0.013 - 0.279
8.05 - 19.75 - 13.43 9.65 26.25 14.97 - 5.85
6.485 12.715 - 2.804 8.752 13.330 12.096 1.763
- 0.400 0.605 - 0.596 - 0.249 - 0.049 - 0.306 - 0.517
-
0.798 0.169 0.191 0.413 0.313 0.124 0.591
2.096 - 2.715 - 1.926 4.429 3.918 3.057 - 0.349
5.73 - 6.97 - 13.45 0.85 - 6.02
- 7.004 - 10.693 - 5.315 4.352 - 8.248
0.842 0.816 - 0.249 - 0.366 0.467
- 0.258 - 0.102 - 0.313 1.598 0.076
-
2.149 1.738 2.349 1.060 1.204
490
L.S. Kaushik et al.: Line • tester analysis for fixed effect model in cotton
Table 6. s.c.a, effects of different crosses for different characters Cross
Yield
Boll no.
Boll wt
No. of monopodes
No. o f s y m p o d e s
1• 4 1• 5 1• 7 1x 8 1• 9
12.27 - 11.10 - 11.82 - 0.70 10.44 23.65
1 • 10
-29.00
1• 1x 1• 1• 1• 2• 2• 2• 2• 2• 2• 2• 2x 2x 2• 2• 2• 3x 3• 3• 3x 3• 3• 3• 3• 3• 3• 3• 3•
-6.71 13.02 - 7.63 5.80 1.74 4.33 - 2.47 2.53 1.50 - 3.77 26.48 -2.00 - 15.28 - 2.45 - 6.90 4.20 -6.26 - 16.62 13.55 9.26 - 0.82 -6.68 - 50.14 30.98 22.00 - 10.56 14.51 - 10.02 4.52
3.493 - 2.707 - 6.185 - 4.507 10.281 15.915 - 14.418 - 7.585 - 0.062 - 2.040 7.226 0.593 4.521 - 1.646 0.676 0.954 5.109 11.310 - 2.424 - 3.657 1.565 - 9.146 - 3.980 - 3.280 - 8.013 4.384 5.509 3.553 - 15.391 - 27.224 16.842 11.243 - 1.502 11.187 - 3.247 2.687
-0.067 0.131 -0.455 0.326 -0.172 -0.128 -0.194 0.254 0.483 - 0.212 -0.191 0.202 0.140 0.028 0.126 -0.038 -0.281 0.279 0.030 -0.182 - 0.356 0.152 0.289 -0.187 -0.073 -0.158 0.330 - 0.314 0.453 -0.150 0.164 - 0.071 -0.125 0.060 - 0.097 - 0.013
- 0.104 0.163 - 0.415 - 0.326 -0.126 0.185 0.418 0.519 0.029 - 0.260 -0.071 - 0.015 0.263 -0.570 -0.098 0.108 0.308 -0.219 -0.181 -0.381 0.163 0.274 - 0.203 0.052 -0.073 0.408 0.463 0.219 -0.181 -0.404 - 0.237 - 0.136 -0.192 -0.015 0.247 - 0.037
3.796 0.207 - 0.949 - 1.071 2.440 3.074 - 4.193 -0.259 - 1.504 0.441 - 0.982 - 1.004 1.102 - 1.421 - 1.510 2.069 0.113 3.147 - 2.120 - 0.287 1.002 0.113 - 0.942 - 1.265 - 0.159 1.212 2.456 - 0.999 - 2.554 -6.221 6.313 0.546 0.502 - 0.554 1.924 2.268
1•
6
11 12 13 14 15 4 5 6 7 8 9 10 11 12 13 14 15 4 5 6 7 8 9 10 11 12 13 14 15
tions. It is i n t e r e s t i n g to n o t e t h a t all five crosses s h o w i n g h i g h e r s.c.a, effects for y i e l d o f s e e d c o t t o n also r e c o r d e d t h e g r e a t e r v a l u e s o f s.c.a, effects for b o l l n u m b e r . T h e s.c.a, for y i e l d o f s e e d c o t t o n a n d boll n u m b e r h a d a l m o s t a p a r a l l e l t r e n d . S i x t e e n cross c o m b i n a t i o n s e x h i b i t e d p o s i t i v e e s t i m a t e s o f s . c . a , effects for boll w e i g h t , t h e m a x i m u m b e i n g in case o f H 7 7 7 x C o c k e r 310, (0.483) f o l l o w e d b y H 8 4 2 x P L 8 0 7 (0.453), H842 • BP68 (0.33), H777 • L a x m i 7882 (0.326). E i g h t e e n c r o s s e s p o s s e s s e d p o s i t i v e s.c.a, effects for n u m b e r o f s y m p o d e s , H 8 4 2 x 152F h a v i n g the m a x i m u m s.c.a. v a l u e (6.313) c l o s e l y f o l l o w e d b y H777 x B A R SP84 (3.796), H655 x H807 (3.14) a n d H777 • H807 (3.074). H y b r i d H842 x 152F, p o s s e s s i n g t h e h i g h e s t s.c.a, effect for n u m b e r o f s y m p o d i a , also p o s s e s s e d t h e h i g h e s t s.c.a, effect for s e e d c o t t o n yield.
Heterotic effects H e t e r o t i c effects w e r e e s t i m a t e d as the d e v i a t i o n s o f the Fl's from m i d - p a r e n t a l values (MP) and the deviations
f r o m its b e t t e r p a r e n t (BP), w i t h r e s p e c t to five traits o f A m e r i c a n cotton. S i x t e e n crosses m a n i f e s t e d h e t e r o s i s o v e r t h e m i d p a r e n t for s e e d c o t t o n yield. C r o s s H842 x H807 (69.81) f o l l o w e d b y H 7 7 7 X P L 8 0 7 (61.81) a n d H 8 4 2 • (61.57) e x h i b i t e d e v e n m o r e t h a n 50% h e t e r o s i s o v e r the mid-parent. Only eight hybrids showed positive h e t e r o s i s o v e r the b e t t e r p a r e n t for s e e d c o t t o n yield, T w o cross c o m b i n a t i o n s , H 7 7 7 • (42.9%) a n d H777 X PL807 (40.2), r e c o r d e d m o r e t h a n 40% h e t e r o s i s o v e r t h e b e t t e r p a r e n t (H777), t h e m o s t c o m m o n l y c u l t i v a t e d c o t t o n v a r i e t y o f the z o n e . Similarly, 16 crosses e x h i b i t e d p o s i t i v e h y b r i d v i g o u r ( M P ) for n u m b e r o f bolls, r a n g i n g b e t w e e n 8.0 to 85.2 (H655 x H807) p e r cent. T w e l v e crosses r e c o r d e d h e t e r o s i s o v e r BP for boll n u m b e r . H y b r i d s H655 x H807 (74.0%), H 6 5 5 • (53.5%) a n d H 6 5 5 • B A R SP (51.4%), r e c o r d e d m o r e t h a n 50% h e t e r o s i s o v e r BP for boll n u m b e r . E l e v e n crosses w e r e f o u n d to b e s u p e r i o r for boll w e i g h t o v e r t h e i r b e t t e r p a r e n t s .
L. S. Kaushik et al.: Line • tester analysis for fixed effect model in cotton Heaviest boll weight was p r o d u c e d b y the crosses H777 x Laxmi 7882 (16.5%), followed by H842 X PL807 (11.9%) and H 7 7 7 x H 8 0 7 (10.5%). More than 50% h y b r i d vigour (BP) was recorded for n u m b e r o f m o n o podes in two crosses, H842 x BP68 (52.2%) and H842 x Reba-B50 (50.8%). Only six crosses recorded h i g h e r n u m b e r o f s y m p o d i a (BP), ranging between 3.0% (H655 X H807) to 28.0% (H777 X H807). The degree of heterosis varied considerably for different characters with the maximum heterosis for the number of bolls per plant, yield of seed cotton and boll weight. Therefore, there appears to be good possibilities of increasing the yield by exploiting hybrid vigour through the number of bolls and boll weight. These results are in confirmity with earlier reports (White and Richmond 1963; Miller and Marani 1963; Marani 1967; Singh and Murty 1971; Singh 1974). For yield of seed cotton, the three best crosses were H842 x 152F, H655XH807 and H777 x H807. These crosses were also best for boll number and the number of both types of branches. Crosses involving parents H842, H807 and 152F exhibited higher degree of heterosis for yield of seed cotton and other component characters of yield in this study. The best combinations (H777xPL807 and H777XH807), as judged from the s.c.a. estimates of seed cotton, involved the best general combiners (Table 6). Therefore, there may be a good chance of success for getting high yielding segregants and it equally may be possible of developing a hybrid involving the parents H777, PL807 and H807. These results are in general agreement with those reported by Turner (1953); Marani (1963, 1964); Gururaja Rao (1975) and Bhallala (1976). In order to exploit the type of gene effects operative in the population developed by involving H777, H807 and PL807, a breeding procedure which takes care of both additive and non-additive gene effects might prove the most efficient in improving the population. Similar findings have also been reported by Gupta and Singh 1970; Singh 1974; Chandermathi and Menon 1973 and Mirza and Khan 1976.
491
References Arunachalam VC (1974) The fallacy behind the use of a modified line x tester design. Indian J Genet 34:280-287 Bhallala MK (1976) Heterosis analysis in upland cotton. MSc Thesis (submitted to HAU, Hisar) Chanderamathi PS, Menon PM (1973) Diallel analysis of yield and its components in G. barbedensi L. Madras Agric J 60: 1560-1566 Fisher RA (1926) The arrangement of field experiments. J Min Agric 33:503-513 Gupta MP, Singh RB (1970) Genetics of some characters in upland cotton. Indian J Genet 30:590-598 Gururajarao MR (1975) Genetic analysis in upland cotton (G. hirstu L.). Mysore J Agric Sci 9:199 Marani A (1963) Heterosis and combining ability for yield and its components in cotton. Crop Sci 3:552-555 Marani A (1964) Heterosis and combining ability in diallel cross of two spps. of cotton. Crop Sci 4:265-268 Marani A (1967) Heterosis and combining ability in intra- and interspecific crosses of cotton. Crop Sci 7:519-522 Miller PA, Marani A (1963) Heterosis and combining ability in varietal top crosses of upland cotton (G. hirsutum L.). Crop Sci 3: 646-649 Mirza AM, Khan MA (1976) Diallel analysis of some important characters in cotton (G. hirsutum L.) Plant Breed (Abstr) 46:3509 Singh BB, Murty BR (1971) Hybrid vigour in intervarietal crosses of upland cotton. Indian J Genet 31:1-7 Singh DP (1974) Genetic architecture of Gossypium hirsutum for certain economic character. PhD Thesis (submitted to Int Adv Study Meerut) Turner JH (1953) A study of heterosis in upland cotton yield of hybrids compared with varieties. Agron J 45:484-486 Yates F (1937) The design and analysis of factorial experiments. Imp Bureau Soil Sci Tech Comm 35:1-95 White TG, Richmond TR (1963) Heterosis and combining ability in top and diallel crosses in cotton. Crop Sci 6:58-63
