Indian Journal of Agricultural Sciences 82 (11): 91621, November 2012

Combining ability and heterosis analysis for yield traits in

bread wheat (Triticum aestivum)
VIKRANT SINGH1, RAM KRISHNA2, SANJAY SINGH3 and PRASHANT VIKRAM4
C S Azad University of Agriculture and Technology, Kanpur, Uttar Pradesh 208 002
Received: 21 December 2011; Revised accepted: 13 July 2012

ABSTRACT
In order to realize the combining ability and heterosis of wheat, 21 hybrids were synthesised in a 7 7 diallel fashion
excluding reciprocals. Analyses of combining ability and heterosis over mid parent (MP) as well as economic parent (HD
2733) were undertaken for yield and its component traits. The experiment was conducted in 200708 and 200809 at
Crop Research Farm of CSAUA &T, Kanpur, UP. The results revealed that non-additive genetic variance played a
predominant role in the inheritance of most of the traits. The best combinations mostly involved high low and low low
general combiners for the characters under study. There was very rare case in which high high general combiners were
involved for best combinations. On the basis of gca and sca effects, 3 parents (i e K 7903, K 9465 and HUW 234) and 14
cross combinations (i.e. 5 top crosses namely HD 2733 K 7903, HUW 234 K 9423, HD 2285 K 2021, HUW 234
K 2021 and K 9423 K 2021) were found good general and specific combiners for higher grain yield and also for various
yield contributing traits, respectively.

Key words: Combining ability, Diallel, Heterosis, Triticum aestivum

methodology to be employed for varietal improvement and

also provides useful information about usefulness of the
parents in breeding programs. So a large number of researches
on heterosis for grain yield and its components in wheat have
been carried out (Singh et al. 2004; Dreisigacker et al. 2005),
since the phenomenon in wheat was first reported in 1919.
Hybrid technology in crop plants, especially cross-pollinated
crops is successfully used for enhanced production. However,
it remains unutilized in the self-pollinated crops, especially
wheat. The future scope of hybrid technology in wheat
depends on the male sterility systems, floral biology, level of
combining ability, heterosis and its exploitation of commercial
level that may be useful in breaking yield barriers and
enhancing the productivity in the major wheat belt of the
country (Singh et al. 2010). Keeping in view of all above, in
the present study, an attempt has been made to examine the
combing ability as well as heterosis of some wheat varieties
and their crosses on yield related traits.

Wheat (Triticum aestivum L. emend Fiori & Paol.) is a

predominant cereal crop of the world and constitutes important
source of carbohydrate and protein. At global level, India
ranks second largest wheat producing nation with 13.43%
global wheat production after China which contributes 17.7%
to the world wheat production (USDA 2012). The other
major wheat producing countries are Russian Federation,
United States of America and Canada and these 5 countries
together contribute more than half of the global wheat
production (Singh et al. 2010). To fulfil the increasing demand
of world population, wheat production and productivity must
be increased. Hybrid wheat is an alternative approach to
increase the productivity and most important step in the
hybrid breeding program is the detection of suitable parents
with high general (gca) and specific combining ability (sca)
for grain yield and then the exploitation of heterosis. The
study of heterosis has a direct bearing on the breeding
1 Research

Scholar (e mail: [email protected]),

(e mail: [email protected]), Department of
Genetic and Plant Breeding, C S Azad University of Agri. and Tech.,
Kanpur 208 002;
3 Senior Scientist (e mail: [email protected]), Seed
Production Unit IARI, New Delhi 110 012
4 Senior Research Fellow (e mail: [email protected]),
Indian Institute of Pulses Research, Kanpur (Present address: Visiting
Research Fellow, International Rice Research Institute, Philippines)
2 Profressor

MATERIALS AND METHODS

Genetic materials, for the present investigation comprised
of seven wheat varieties (K 9465, K 9423, K 2021, K 7903,
HD 2733, HD 2285 and HUW 234) and 21 hybrids generated
by crossing the above varieties in all possible combinations
excluding reciprocals, were evaluated in randomized block
design with 3 replications and the experiment was conducted
8

November 2012] COMBINING ABILITY AND HETEROSIS ANALYSIS FOR YIELD TRAITS IN BREAD WHEAT

in 200708 and 200809 at Crop Research Farm of CSAUA

& T Kanpur, UP. All recommended agronomic practices
were adopted in the trial shown in 3 rows of 3 m length with
inter- and intra- row spacing of 25 cm and 15 cm, respectively.
Five randomly selected competitive plants were observed
for 14 traits namely days to flowering (from date of sowing
to date of 75% flowering stage), days to maturity (from the
date of sowing to its 75% maturity), duration of reproductive
phase (days to maturity - days to flowering.), number of
productive tillers/plant (total numbers of ear bearing tillers
per plant at the time of maturity), plant height (from ground
level to tip of ear avoiding awns of the panicle of the main
shoot at maturity in cm), spike length (from base of spike to
the tip of the ear avoiding awns in cm at maturity), number
of spikelets per spike (the spikelets bearing the grains in each
spike), number of grains per spike (average number of grains
per spike of selected plants), biological yield (weight of
whole selected plants including root in gram), harvest index

917

(%), test weight (gram), seed hardness (by O.S.K. 201 grain
hardness Tester, type E capacity 50 kg in kg/Seed), protein
content (by Micro-Kjeldahl method in %) and grain yield per
plant (gram). The mean values were used for estimating the
heterosis and combining ability according to Griffings (1956,
a) model l method 2, which is a fixed effect model.
RESULTS AND DISCUSSION
Average as well as economic heterosis
Heterosis for all the 14 traits studied was found with all
crosses. A wide variation of heterosis range, heterosis mean,
number of desired hybrids and best hybrid was found for
most of the traits (Table 1). Singh et al. (2004) stated that the
superiority of hybrids particularly over high parent is more
useful for commercial exploitation of heterosis and also
indicated the parental combinations capable of producing
the highest level of transgressive segregants. However, for

Table 1 Heterosis range, heterosis mean, number of desirable hybrids and best hybrid [over economic variety (HD 2733 and mid parent
(MP)] for 14 traits in bread wheat
Trait
Days to flowering
Days to maturity
Duration of reproductive phase
Number of productive tillers/plant
Plant height
Spike length
Number of spikelets/spike
Number of grains/spike
Biological yield
Harvest index
Test weight
Seed hardness
Protein content
Grain yield/plant

Heterosis
hybrids
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP
EV
MP

Heterosis range
(%)
18.87 to 4.21
6.67 to 15.99
31.07 to 0.37
5.91 to 20.65
63.43 to 0.82
20.25 to 32.27
1.97 to 113.18
1.75 to 113.13
4.66 to 48.13
20.85 to 39.91
19.32 to 13.19
14.81 to 33.07
7.18 to 9.20
3.58 to 16.23
24.20 to 28.41
21.6 to 41.26
20.05 to 69.20
22.47 to 105.04
17.32 to 50.23
24.08 to 73.54
4.94 to 49.74
12.16 to 40.39
18.99 to 0.34
14.54 to 21.08
10.10 to 4.22
6.76 to 11.72
1.14 to 79.35
2.58 to 114.64

Heterosis
mean

No. of desired
hybrids

5.0
3.27
9.50
5.52
16.10
8.79
36.48
29.97
10.96
2.64
0.96
10.12
0.11
5.34
1.20
3.82
30.34
31.51
16.35
23.45
20.84
12.08
8.80
3.17
2.77
1.25
51.56
59.22

17
7
20
4

E V, Average value of economic variety (HD 2733); M P, average value of mid parent
9

13
21
17
4
12
10
16
9
15
6
13
19
19
15
17
20
17
11
3
7
19
19

Best hybrids
HD 2285 K 7903
HD 2285 K 7903
K 9423 K 7903
HD 2733 K 9465
K 2021 K 7903
HD 2733 K 9423
HD 2285 K 2021
HD 2285 K 2021
HUW 234 K 2021
HUW 234 K 2021
K 9423 K 9465
K 2021 K 7903
HD 2285 K 7903
HD 22895 K 7903
HD 2285 K 7903
HD 22895 K 7903
HUW 234 K 9465
HUW 234 HD 2285
HUW 234 K 2021
K 9423 K 7903
K 9423 K 9465
HD 2733 HUW 234
K 9423 K 9465
HD 2733 HUW 234
K 9465 K 7903
HD 2285 K 7903
HUW 234 K 9423
HD 2285 K 2021

10

* Significant at 5% level, ** significant at 1% level; gca, general combining ability; sca, specific combining ability; 2g / 2s, ratio of gca variance to sca variance; ( 2s/ 2g )0.5, degree
of dominance

6.55**
16.15**
0.16
0.71
15.99
0.04
4.75
0.62
0.24
0.01
0.07
0.23
0.3
1.81
0.29
0.67
0.01
0.03
0.66
0.05
4.69
17.31** 49.89** 52.39** 19.26**
34.45** 47.06** 97.93** 23.54**
0.14
0.04
0.07
0.02
1.91
5.54
5.81
2.14
34.31
47.02
97.86
23.52
0.06
0.12
0.06
0.09
4.24
2.91
9.11
3.32
1.23
1.13
0.03
0.13
1.1
0.12
2.91
0.63
1.51
0.01
0.07
1.5
0.05
4.63
0.5
36.76**
4.10** 65.89**
0.02
0.04
0.05
4.08
4.08
65.85
0.01
0.06
9.03
4.02
45.85** 477.85** 252.04**
10.06** 43.67** 41.20**
0.03
0.03
0.02
5.09
53.09
28
10.03
43.64
41.18
0.51
1.22
0.68
1.4
0.91
1.21
6
21
54
Gca
Sca
Error
2g
2s
2g / 2s
( 2s/ 2g )0.5

Test
Seed
Protein
weight hardness content
(g)
(kg/seed) (%)
Harvest
index
(%)
Biological
yield
(g)
Days to
flowering

Days to
maturity

Duration
of reproductive
phase
(days)

Number
of productive
tillers/
plant

Plant
height
(cm)

Characters
Spike Number Number
length
of
of
(cm) spikelets/ grains/
spike
spike
df

Combining ability variances and effects

General and specific combining ability variances and
effects were estimated with a view to decipher the genetic
architecture of the characters under study. Combining ability
describes the breeding value of parental lines to produce
hybrids (Romanus et al. 2008). The general combining ability
has been equated with additive gene action and specific
combining ability with non-additive gene action (Griffing
1956 a). The analysis of variance for combining ability was
done for all the 14 characters (Table 2). Highly significant
variances, of both general and specific combining ability,
were observed which indicated the importance of both additive
and non-additive gene effects for all the traits except number
of productive tillers per plant (where only sca variance was
highly significant indicating non-additive gene action), spike
length, number of spikelets per spike, seed hardness and
protein content. The estimated value of 2g was higher than
its 2s for days to maturity which indicated the predominance
of additive gene action as the ratio of 2g/ 2s was more than
unity while rest of the traits showed preponderance of nonadditive gene action. The value of average degree of
dominance ( 2s/ 2g)0.5 for days to maturity indicated partial
dominance while rest of the traits showed over dominance.
In the same way, preponderance of non-additive gene effects
were reported by several researchers (Singh 2003, Chaman
2005, Heidari et al. 2006 and Kumar et al. 2011) for plant
height, number of tillers per plant, spike length, number of
spikelets per spike, test weight, seed hardness, protein content
and grain yield.
Selection of suitable genotypes and their crosses in
effective hybridization is a pre requisite in order to formulate
a systematic breeding programme leading to rapid and
sustained improvement. The combining ability effects (Tables
3, 4) furnish information on these aspects. Thus, the genetic
material available in the present study is suitable for evolving
desirable hybrids as well as varieties. In the later case, some
sort of intermating within segregating progenies at various
stages would be more desirable to harness additive and
additive additive type of variance.
While considering gca effects of the parents, it was

Sources of variation

grain yield per plant, economic and average heterosis ranged

from 1.14 to 79.35 and from 2.58 to 114.64 with mean value
of 51.56 and 59.22, respectively. Early flowering, early
maturity and short stature are the desirable traits in wheat.
HD 2285 K 7903 and HUW 234 K 2021 showed highest
negative heterosis for days to flowering and plant height,
respectively both over economic variety and mid parent.
K 9423 K 7903 and HD 2733 K 9465 showed highest
negative heterosis for days to maturity over economic variety
and mid parent, respectively. Significantly highest positive
economic and average heterosis was observed in HUW 234
K 9423 and HD 2285 K 2021 for grain yield/plant,
respectively.

[Indian Journal of Agricultural Sciences 82 (11)

Grain
yield
(g)

SINGH ET AL.
Table 2 Analysis of variance (ANOVA) for combining ability along with estimates of genetic components of variance and degree of dominance for 14 characters in bread wheat

918

11

I, VI, IV
I, III
III, II
I, V
VII, IV
VII, IV
C.P.

A = SE (gi) +, B = SE (gi-gj) +, G.P. = good general combiner parents, H.P. = High per se performance parents, C.P. = Common parents based on gca and per se performance

I, IV

I, IV
I, VI, IV
I, III
III, II
I, V
VII, IV
VII, IV
H.P.

* Significant at 5% level, ** significant at 1% level, I = HD 2733, II = HUW 234, III = HD 2285, IV = K 9423, V = K 2021, VI = K 9465, VII = K 7903

VI, VII, I

VII, VI, II

0.19

VI, VII
V, VII
III, IV
VII, VI, IV
II, I
VII, VI
IV, I

II, III, VI, I VII, VI, V, IV I, III, VII, IV IV, V, I, VII

VII, VI
IV, I

VII, V
III , IV, VI
VII, VI, IV
II, IV, I
I, VI, IV
II, I, III
II , III
VII, IV
G.P.

0.09

0.06

IV, VII

0.07

0.05

0.07
0.08
B

0.05
0.05
A

V, I, VI, III, II

0.06
0.03

0.04

0.09

IV, I

0.12

0.18

IV, I

VI, VII, III

0.12
0.04

0.05
0.06

0.04
0.04

0.07
0.12

0.08
0.06

0.09

919

0.1

0.06

11.7 0.62** 14.2

-0.06
9.5

39.6 2.36** 39.7 -0.10* 10.8 0.32** 12.3 0.90** 13.9

18.5 -0.32* 50.3 2.36** 35.5 0.12
-0.14

0.27** 62.3 0.25** 11.0 -0.23** 19.5 -1.63** 57.1 3.28** 30.5 -3.16** 46.6 0.90** 37.0 0.12*

0.55** 64.1 -0.29** 8.2

7.9
VII

116.1 1.89** 48.5 -0.13* 7.1

0.28** 67.9 2.08**

-4.10** 54.5 -10.00** 83.1 -6.14** 29.3 -0.12

VI

10.9
12.8 -0.03
10.3 0.08

10.1 0.23** 12.6 -0.46** 8.9

27.2 -2.55** 32.2 -2.15** 36.0 0
-0.43** 18.3 -1.24** 53.7 -0.05

10.3 0.54** 19.9 2.00** 64.1 -1.12** 26.7 1.57** 40.8 0.78** 35.0 0.12*

0.92** 79.9 -0.23** 8.8

3.12** 68.9 0.08*

7.3

122.9 5.67** 56.6 -0.30** 6.3

-1.36** 62.7 -10.02** 84.4 -8.47** 23.1 0.04

2.03** 67.1 7.63**

IV

9.7

9.7
11.4 -0.45** 10.7 0.2
56.9 0.51** 19.4 -0.26* 50.2 -0.63** 34.8 0.14*
19.7 0.05
-0.36** 68.4 -0.13** 10.1 0.02
69.5 1.96**
0.04
III

117.9 1.72** 49.2 0.41** 6.9

12.6 -1.65** 11.7

11.9 0.07

-0.23** 19.1 -0.57** 52.5 -1.27** 17.7 3.98** 54.7 -0.45** 34.8 -0.36** 10.1 -0.19** 11.9 0.43*

-0.97** 62.8 0.43** 10.9 0.47** 20.3 1.71** 58.9 -3.71** 25.1 0.31** 46.5 -0.80** 33.2 0.08

-3.54** 71.9 -0.11* 9.5

6.6

6.6
127.7 4.34** 57.1 -0.08

II

113.5 0.99** 47.5 0.18*

2.76** 71.3 7.09**

0.34** 66.7 1.26**

Per
se
gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Per
se

gca

Protein
content
(%)
Seed
hardness
(kg/seed)
Test
weight
(g)
Harvest
index
(%)
Biological
yield
(g)
Number of
grains/
spike
Number of
spikelets/
spike
Spike
length
(cm)
Plant
height
(cm)
Number of
productive
tillers/plant
Duration of
reproductive
phase (days)
Days
to
maturity
Days
to
flowering
Parents

Table 3 Estimates of gca effects and per se performance as well as their ranking (only desired)of the parents for 14 characters in bread wheat

gca

Grain
yield
(g)

found that none of the parents was observed as good general

combiner for all the 14 traits. However, K 7903 and K 9423
for early flowering and early maturity, K 2021 and HD 2733
for longer duration of reproductive phase, HUW 234 and
HD 2285 for more number of productive tillers per plant,
HUW 234, HD 2733 and HD 2285 for short stature, HD
2733, K 9465 and K 9423 for lengthy spikes, HD 2733 and
K 9423 for more number of spikelets per spike, K 9423 and
HD 2733 for more number of grains per spike, K 7903 and
K 9465 for more biomass, HUW 234, K 9423 and HD 2733
for high harvest index, K 7903, K 9465 and K 9423 for
bolder seed size, HD 2285 and K 9423 for hard seed, K 7903
and K 2021 for high protein content and K 7903, K 9465 and
HUW 234 for high grain yield were G.P. (as in descending
order for each characters, Table 3). Further, the comparison
of per se performance with gca effects of the genotype
provide a wholesome view of its breeding worth. In the
present study, it was noticed that there was close relationship
between parental mean performance and gca effects for
almost all the characters. Hence, these parents can be
employed in breeding programmes for overall improvement.
Among the 21 hybrids, only 14 cross combinations were
found significant and desirable for higher grain yield and
also for other traits (Table 4). On the basis of sca effects and
per se performance of the hybrids, it was noted that the
crosses were not exactly in the same order of ranking. In the
present findings, best combinations mostly involved high
low and low low general combiners for the characters
under study. There was very rare case in which high high
general combiners were involved for best combinations (Table
4). The same type of result was also observed by Kumar and
Maloo (2012). Thus, it is evident that high specific combiners
are not always obtained between high general combiners but
may occur between low low or high low general
combiners. This might be probably due to the presence of
dominant and epistatic gene interactions.
In general, sca effects do not make any significant
contribution in the improvement of self-pollinated crops
except where there is possibility of commercial exploitation
of heterosis. Breeders interest normally, however, vests in
obtaining transgressive segregants through crosses in order
to produce homozygous lines in self-pollinated crops.
Therefore, crosses involving high low general combiners
in respect of different characters in the present study may be
utilized for obtaining transgressive segregants in the next
generation resulting from dominance gene interaction. Thus
based on gca, the varieties K 7903, K 9465 and HUW 234
could be a better choice for the improvement of yield and
component traits through hybridization. The crosses HD 2733
K 7903, HUW 234 K 9423, HD 2285 K 2021,
HUW 234 K 2021, K 9423 K 2021, HD 2285 K 9465,
HUW 234 HD 2285, K 9423 K 9465, HD 2285 K 9423,
HD 2733 HUW 234, K 2021 K 9465, K 2021 K 7903,
HD 2733 HD 2285 and HUW 234 K 7903 which had

Per
se

November 2012] COMBINING ABILITY AND HETEROSIS ANALYSIS FOR YIELD TRAITS IN BREAD WHEAT

sca

sca

sca

Per
se

sca

Per
se

sca

Per sca
se

26.8 7.49**

3.31** 69.2 -2.84** 107.8 -6.16** 39.1 -0.42** 7.9

-1.74** 65.9 -2.38** 114.4 -0.74** 49.0 -2.09** 7.0

1.06** 67.3 -1.07** 103.7 -2.15** 37.4 3.69** 12.4 -7.90** 61.0 0.67** 11.2 0.53** 20.9 3.26** 62.4 6.62** 35.8 1.24**

4.74** 74.3 1.82**

II x III

II x IV

II x V

12

2.12** 68.3 7.85**

6.16** 67.9 -5.53** 88.0 -11.55** 20.9 -0.94** 7.5

-0.67** 68.9 -6.06** 117.2 -4.99** 49.6 -1.33** 6.7

3.11** 68.3 13.09** 124.3 10.04** 56.6 -0.80** 7.3

1.06** 64.5 9.04**

IV x VI

IV x VII

V x VI

V x VII

VI x VII

0.25

0.17

9.1

0.2

0.13

114.7 8.10**

113.5 5.42**

113.7 5.84**

108.5 6.93**

117.5 0.23

8.2

8.4

0.21

14

50.9 062** 8.9

45.9 -0.2

50.1 0.18

49.5 0.88** 9.7

50.9 -0.91** 7.9

Per
se

sca

Per
se

59.3 -2.16** 36.8 -0.17

10.8 0.18

12.5 -2.50** 11.9

37.8 -0.76** 10.3 -0.23* 11.6 1.20** 15.8

0.61** 12.6 2.14** 17.0

sca

57.1 -3.08** 28.1 -1.77** 47.1 0.33* 39.4 0.09

-0.1

12.5 5.33** 20.7

11.1 -0.25* 11.9 -1.78** 13.3

59.8 1.91** 32.1 12.21** 64.3 -0.83** 39.7 -1.09** 9.7

58.5 -0.57** 38.8 0.14

10.7 -0.57** 11.5 4.46** 20.9

-8.24** 63.9 -1.21** 9.3

-4.81** 61.6 -1.03** 9.1

19.5 0.64

57.5 -0.73** 31.9 2.21**

40.7 0.02

17.3

0.14

10.9 -0.1

12.1 4.21** 20.0

11.7 2.22** 18.5

10.4 -0.55** 11.8 1.05** 18.5

49.3 -2.22** 34.0 -1.19** 9.8

54.5 4.31** 43.5 -0.2

58.0 -0.21

73.7 -0.31** 9.8

0.87** 21.1 -0.21

58.3 1.86** 32.3 8.43**

16.9
59.1 -2.37** 35.3 0.87** 11.8 0.24* 12.7 3.48** 19.1

2.22** 22.2 18.21** 75.7 3.02** 37.5 -1.75** 49.8 -2.66** 38.1 -0.35** 10.5 0.82** 12.9 -0.32

10.7 -0.05
55.6 0.49** 38.2 0.60** 11.5 -0.09

12.5 0.25

17.2
12.3 1.89** 18.1

0.27

0.18

4.15**
0.11

0.08
0.25

0.17

74.3 0.48** 11.0 -0.27

0.51

0.34

19.5 0.49

0.27

0.18

0.35

0.24

0.19

0.13

0.16

0.11

0.15

0.1

56.3 2.45** 39.7 -3.38** 45.3 6.48** 48.7 0.48** 11.3 0.66** 13.1 0.36

0.53

0.36

18.0

-5.83** 65.0 1.26** 11.3 -0.39* 19.1 -3.90** 52.3 3.18** 37.1 -0.93** 48.3 2.23** 41.4 0.69** 11.4 0.27* 13.0 1.38** 17.9

-2.29** 68.3 1.42** 12.0 -0.44* 19.0 3.35** 58.2 -2.14** 32.7 9.59**

20.04** 93.1 1.75** 12.1 0.50** 21.0 -2.47** 56.9 -8.15** 24.7 16.36** 69.7 3.02** 45.2 -0.09

10.22** 83.0 1.44** 12.3 1.53** 21.9 4.84** 62.9 8.73** 42.5 -5.19** 44.9 8.98** 49.7 0.92** 12.0 -0.63** 11.6 2.40** 19.1

-0.70** 72.7 -1.61** 8.8

4.15**

-5.60** 63.7 -0.58** 10.1 -1.02** 18.9 -11.48**44.7 -2.10** 33.3 12.27** 60.5 -0.70** 38.6 0.83** 11.9 0.70** 12.4 3.16** 20.1

57.0 8.63** 40.7 0.39

-0.39* 20.3 -9.10** 50.7 2.97** 33.9 1.47**

-0.26

0.90** 20.5 -1.85** 53.7 11.48** 45.1 -14.04** 38.5 -7.92** 31.5 0.24* 10.8 0.40** 12.3 0.21

10.2 -0.56** 18.9 2.69** 58.6 -2.05** 28.2 16.78** 69.9 -1.64** 34.8 -0.48** 10.0 0.24* 12.5 3.65** 19.7

I = HD 2733, II = HUW 234, III = HD 2285, IV = K 9423, V = K 2021, VI = K 9465, VII = K 7903,
A = SE (gi) +, B = SE (gi-gj) +, * Significant at 5% level, ** significant at 1% level

0.21

-3.43** 64.5 2.50**

IV x V

-5.31** 57.9 2.96**

III x VII

0.14

-0.28

III x VI

43.9 0.12

Per
se

72.2 1.42** 11.7 -0.88** 19.0 1.14** 58.3 7.19** 38.0 -3.80** 51.6 6.91** 77.8 0.98** 11.6 -0.24** 11.3 3.02** 19.7

-6.84** 59.9 0.05

6.71**

123.7 -1.82** 52.6 5.46** 14.1 -2.22** 67.7 0.94** 11.1 1.65** 21.3 0.47

107.3 3.59**

20.0 -0.68

sca

Grain
yield
(g)

SINGH ET AL.

2.44** 71.7 0.53**

67.3 -0.13

-1.85** 64.1 1.77**

III x V

-0.34

-4.98** 64.0 1.43** 12.1 0.77** 21.2 0.68

51.3 2.12** 10.7 -2.43** 63.7 -0.73** 9.9

45.5 1.31** 9.9

III x IV

109.1 3.59**

117.5 1.36**

-0.25

0.80** 64.3 4.25**

II x VII

124.3 -3.22** 50.5 -0.77** 7.6

II x VI

67.6 0.64**

-2.27** 68.0 -8.05** 114.7 -5.59** 47.7 2.05** 10.3 11.20** 79.9 -1.91** 9.3

Per
se

Protein
content
(%)

58.8 -0.95** 26.9 -5.55** 43.9 0.91** 37.0 0.87** 11.8 -0.71** 11.8 -2.17** 11.8

71.4 -0.20* 10.8 -0.44* 20.7 -1.03** 60.4 -6.68** 20.1 5.74**
69.7 0.54** 11.3 0.67** 20.8 0.61

I x VII

0.42*

sca

Seed
hardness
(kg/seed)

63.5 9.98** 47.7 -0.86** 9.7

Per
se

Test
weight
(g)

10.8 -0.65** 19.9 2.65** 62.1 6.69** 35.1 -6.57** 45.2 0.23

I x VI

128.1 -2.31** 54.7 0.49** 8.6

-0.11

73.7 0.01

Harvest
index
(%)

Per sca
se

Biological
yield
(g)

-0.36* 68.3 12.71** 123.3 12.90** 55.8 -0.25

8.2

Per
se

Number of
grains/
spike

-3.02** 61.8 1.16** 12.0 1.40** 21.7 2.21** 61.1 0.21

sca

Number of
spikelets/
spike

2.12** 74.1 -0.13

8.9

Per
se

Spike
length
(cm)

IxV

56.5 0.27

Per
se

Plant
height
(cm)

I x IV

123.9 4.10**

Per
se

Number of
productive
tillers/plant

2.30** 72.3 -3.53** 119.1 -5.63** 47.5 1.31** 10.1 5.67**

sca

Duration of
reproductive
phase (days)

-2.26** 68.1 1.96**

Per
se

Days
to
maturity

Ix III

sca

Days
to
flowering

I x II

F1s

Table 4 Estimates of sca effects and per se performance of 21 F1s for 14 characters in bread wheat

920
[Indian Journal of Agricultural Sciences 82 (11)

November 2012] COMBINING ABILITY AND HETEROSIS ANALYSIS FOR YIELD TRAITS IN BREAD WHEAT

921

Heidari B, Rezai A and Maibody S A M M. 2006. Diallel analysis

for the estimation of genetic parameters for grain yield and grain
yield components in bread wheat. Journal of Science and
Technology of Agriculture and Natural Resources 10 (2): 121
40.
Kumar A, Mishra V, Vyas R P and Singh V. 2011. Heterosis and
combining ability analysis in bread wheat (Triticum aestivum
L.). Journal of Plant Breeding and Crop Science 3(10): 20917.
Kumar V and Maloo S R. 2012. Parental molecular diversity and its
concurrence to heterosis in bread wheat (Triticum aestivum).
Indian Journal of Agricultural Sciences 82 (3): 20712.
Romanus K G, Hussein S and Mashela W P. 2008. Combining
ability analysis and association of yield and yield components
among selected cowpea lines. Euphytica 162: 20510.
Singh H, Sharma S N and Sain R S. 2004. Heterosis studies for
yield and its components in bread wheat over environments.
Hereditas 141: 10614.
Singh S K, Chatrath R and Mishra B. 2010. Perspective of hybrid
wheat research: A review. Indian Journal of Agricultural Sciences
80 (12): 1 01327.
Singh S K. 2003. Gene action and combining ability in relation to
development of hybrids in wheat. Farm Science Journal 12 (2):
11821.
USDA. 2012. Grain: World Markets and Trade, May 2012,
www.fas.usda.gov/psdonline, 10 pp.

highly significant sca effect for most of the yield contributing

traits may be exploited for the development of single cross
hybrids and also through the population improvement
programme for the development of suitable high-yielding
varieties after knowing the extent of depression which
inbreeding could reign in the F2 and subsequent generations.
ACKNOWLWDGMENTS
Studentship support, provided by the Chandra Shekhar
Azad University of Agriculture & Technology to conduct
present work as part of M Sc thesis, is duly acknowledged.
REFERENCES
Chaman S, Gupta S K and Satija D R. 2005. Genetic architecture
for some quality traits in wheat (T. aestivum L.). Indian Journal
of Genetics and Plant breeding 65 (4): 27880.
Dreisigacker S, Melchinger A E, Zhang P, Ammar K, Flachenecker
C, Hoisington D and Warburton M L. 2005. Hybrid performance
and heterosis in spring bread wheat, and their relations to SSRbased genetic distances and coefficients of parentage. Euphytica
144: 519.
Griffing B. 1956 a. Concepts of specific and general combining
ability in relation to diallel crossing systems. Australian Journal
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