International Journal of Basic & Applied Sciences IJBAS-IJENS Vol:14 No:01

Combining Ability and Heritability Studies for

Important Traits in F2 of Brassica Napus
*Ali Muhammad, Raziuddin, Ashiq Muhammad, Haneef Raza, Aziz Ur Rahman and Imtiaz Ali
*Author for Correspondence: alikhaksar87@gmail.com
Department of Plant Breeding and Genetics
The University of Agriculture, Peshawar-Pakistan
Abstract--

The present study was undertaken to estimate

combining ability and heritability in 4 4 diallel crosses of
Brassica napus L. at the University of Agriculture, Peshawar
during 2012-13 crop season. The genetic material comprised
four parental genotypes along with their 12 F2 populations grown
in randomized complete block design with three replications.
Analysis of variance revealed significant (P0.01) differences for
plant height, main raceme length, pod length, while significant
(P0.05) differences were observed for days to 50% flowering.
Parental genotype G6 was the best general combiner for days to
50% flowering, plant height, pod length. G9 had the highest
GCA for main raceme length. Cross G2 G4 was the best
specific combiner for plant height, pod length. For reciprocal
effects cross G6 G4 was a good combination with highest
reciprocal effects for main raceme length. Broad sense
heritability recorded was 0.26, 0.52, 0.65 and 0.73 for days to
50% flowering, main raceme length, plant height and pod length
respectively, suggesting the effectiveness of selection of these
traits in different generations. The overall study showed the
importance of both additive and non-additive gene actions.

Index Term-

Brassica napus, combining ability, GCA, SCA,

RCA, heritability and F2 population

INTRODUCTION
Brassica napus is the major crop of genus Brassica and
family Brassicaceae or cruciferae. It is commonly known as
rapeseed, oilseed rape or canola is used as major vegetable
oil source. Brassica seed oil is considered as the major
source of edible oil in Indo-Pak subcontinent Pakistan. At
present, total edible oil requirement in the country is 3.079
million tons of which 0.696 million tons contributed by
local production (34 percent of the requirement); while
remaining requirement was met through import of 2.383
million ton. The import bill cost Rs. 224 billion (US$ 2.611
billion) in 2010-11 (PBS, 2012). In Pakistan, during 20102011, brassica was planted on 575 thousand acres which
produced 203 thousand tons seed with oil production of 61
thousand tons (PBS, 2012).

In rapeseed breeding program for hybrid and open

pollinated varieties, general and specific combining ability
effects (GCA and SCA) are important indicators of the
potential of inbred lines in hybrid combinations. To
incorporate desirable characters to maximize economic
yields, the knowledge of combining ability is valuable to get
information on selection of parents and nature of gene
actions involved. This important information could provide
an essential tool for the rapeseed breeders in the selection of
better parental combination for further improvement (Sher
Aslam Khan et al, 2009; Panhawar et al, 2008). General
combining ability is used to specify the average
performance of a line in hybrid combination. Specific
combining ability is used to indicate those cases in which
certain combinations do relatively better or worse than
would be expected on the basis of the average performance
of the lines involved (Sprague and Tatum 1942). Most of
the quantitative characters are controlled by polygene,
which are influenced by the environment. Hence, it is
essential to partition the overall variability into heritable and
non-heritable components with the help of heritability. The
objectives of the present study were to identify good
combiners for future breeding and to estimate heritability
for various traits in segregating generations of Brassica
napus.
MATERIALS AND METHODS
This experiment was conducted to determine combining
ability and heritability for important traits in F2 populations
of Brassica napus L. at the research farm of Department of
Plant Breeding and Genetics, The University of Agriculture,
Peshawar during 2012-13. Genetic material comprised a set
of four Brassica napus L. genotypes viz., G2, G4, G6, G9
and their 12 F2 populations (G2 G4, G2 G6, G2 G9,
G4 G2, G4 G6, G4 G9, G6 G2, G6 G4, G6 G9,
G9 G2, G9 G4, G9 G6). The parental genotypes were
provided by China and their F1 populations were developed
in The University of Agriculture, Peshawar in 2011-12.

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International Journal of Basic & Applied Sciences IJBAS-IJENS Vol:14 No:01

Table I
List of genotypes used in the study

Genotypes Codes

Source

G2

DH-1136-4-4

G4

DH-1136-1-117

G6

DH-1147-2-26

G9

DH-1209-15-67
RESULTS AND DISCUSSION

All the parental genotypes along with their F2 populations

were sown in randomized complete block design with three
replications. Row to row distance was 60 cm while plant to
plant distance was 20 cm and row length was 4 m. Data
were recorded on the following parameters on 15 randomly
selected plants in each genotype for days to 50% flowering,
plant height, main raceme length and pod length.
Statistical analysis
Analysis of variance
The data taken were statistically analyzed according to the
appropriate method suggested for randomized complete
block design (Steel and Tori, 1980), while mean separation
was carried out through LSD test at 5% level of probability.
The variance due to crosses will be partitioned into the
variance due to general combining ability effects, specific
combining ability effects and reciprocal effects by the
following formula for Method I of model I (Griffings,
1956).
General combining ability (GCA)

gi =

1
1
( Y i . + Y . i) - 2 Y..
2n
n

Specific combining ability (SCA)

sij =

1
1
1
( Y ij +Y ji ) ( Y i .+Y . i +Y j .+Y . j) + 2 Y..
2
2n
n

Reciprocal effects (REC)

r ij =

1
( Y ij - Y ji )
2

Heritability (B.S)
Variances components and heritability estimates were
calculated by the following formula suggested by Singh and
Chowdhury (1985);

( BS )

g
p

2g
= 2
p

Days to 50% flowering

In brassica early flowering results in comparatively larger
grain filling period, which finally results into bold grains
hence negative GCA and SCA effects are preferred.
The mean squares showed that the effects due to SCA were
significant (P0.05) while non-significant (P>0.05)
differences were observed for GCA and RCA (Table II).
These results revealed the presence of non-additive gene
actions in the expression of days to 50% flowering. Nonsignificant RCA indicated the absence of cytoplasmic genes
contributed by the female parents. Among parents G2 was
the best general combiner as it indicated the desired
minimum GCA (-0.78) for days to 50% flowering followed
by G6 (-0.16) as presented in the Table III.
Among the F2 populations, G4 G9 and G6 G9 were the
best specific combinations, as it revealed the desired
minimum SCA (-2.17) followed by G2 G6 (-1.58) as
shown in the Table IV. The desired minimum reciprocal
effect was indicated by the cross G9 G6 (-0.62) followed
by G9 G2 (-0.1) as showed in the Table V. Our results
suggested the presence of both types of gene action for days
to 50% flowering. Huang et al. (2009) and Oghan et al.
(2009) also reported significant GCA and SCA effects in
Brassica napus. Recently, Maurya et al. (2012) also
reported similar results in Brassica juncea.
Genetic and environmental variances for days to 50%
flowering were 4.66 and 13.08 respectively, while
heritability (bs) was 0.26 (Table VI). Marjanovic et al.
(2011) also reported low heritability for days to 50%
flowering in Brassica napus. Similarly, Oghan et al. (2009)
reported high heritability for days to 50% flowering in
Brassica napus L. while Patel and Vyas (2011) reported
moderate to high heritability for days to 50% flowering in
Brassica juncea L..
Plant height (cm)

= Genotypic variance for a trait.

= Phenotypic variance for a trait.

(BS) = Broad sense heritability for a trait.

As shown in the Table II, both GCA and SCA effects were
highly significant (P0.01), while RCA showed nonsignificant (P>0.05) differences. Non-significant RCA
effects indicated the absence of cytoplasmic genes given by
the female parents. Taller plants mostly face the problem of
lodging therefore short stature plants were appropriate in
brassica hence negative GCA and SCA were preferred.
Among parental genotypes, G2 indicated the desired

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International Journal of Basic & Applied Sciences IJBAS-IJENS Vol:14 No:01

minimum GCA (-4.98) and was the best general combiner
for plant height tracked by G4 (-1.92) as indicated in the
Table III. Our results are further strengthened by Rameeh
(2012) who reported the presence of both additive and nonadditive gene actions in the appearance of plant height
showing that selection would bring about significant
improvement in Brassica napus L..
G2 G9 was proved as the best specific combiner among
the F2 populations, as it acquired the desired minimum SCA
(-1.51) as presented in the Table IV. The desired minimum
reciprocal effect was indicated by the combination G9 G2
(-0.88), as observed in Table V. The present findings are in
agreement with the findings of Akbar et al. (2008) and
Azizinia (2011) they found significant GCA and SCA
effects for plant height and revealed the importance of both
types of gene actions in Brassica napus L.. Dar et al. (2013)
observed significant GCA effects in Brassica rapa L.
indicated the presence of additive gene action for plant
height. Rameeh (2011) found significant SCA effects for
plant height showed the presence of non-additive gene
action in Brassica napus L..
Genetic and environmental variances for plant height were
97.68 and 53.5 respectively, while heritability (bs) was high
as 0.65 (Table VI). Our results are in agreement with the
findings of Shehzad and Farhatullah (2012). They found
high heritability in F2 population of Brassica napus. High
heritability estimates have also been reported earlier by
Mahak et al. (2011) in Indian mustard. However, Ali et al.
(2002) and Ali et al. (2003) reported low broad sense
heritability for plant height in Brassica napus L..
Main raceme length (cm)
Combining ability analysis showed highly significant
(P0.01) differences for GCA and SCA (Table II). Among
Parents, G9 acquired maximum GCA (2.58) and was
considered the best general combiner for main raceme
length followed by G6 (1.95), as indicated in the Table III.
Similar results were also reported by Singh et al. (2011) in
Indian mustard (Brassica juncea) and found significant
GCA and SCA effects for main raceme length. These
outcomes proposed that both additive and non-additive gene
actions were complicated in controlling main raceme length.
Among the F2 descendants, G2 G6 was the best specific
combiner, as it exhibited the maximum SCA (3.9) followed
by G4 G6 (2.25), as showed in the Table IV. The maternal
effects were non-significant (P>0.05) for main raceme
length due to the lack of cytoplasmic genes given by the

female parents. Cross combination G6 G4 exhibited the

desire maximum RCA (5.3) followed by G4 G2 (2.1) and
G9 G6 (0.85), as indicated in the Table V. Similar results
were earlier reported by Gupta et al. (2011) and Singh et al.
(2010) they found significant GCA and SCA effects for
main raceme length in Brassica juncea.
Genetic and environmental variances for main raceme
length were 22.35 and 20.91 respectively, while heritability
(bs) was 0.52 (Table VI). Nasim et al. (2013) reported low
heritability for main raceme length in Brassica napus L. Ali
et al. (2013) reported high heritability for main raceme
length in Brassica carinata.
Pod length (cm)
Combining ability analysis showed highly significant
(P0.01) differences for GCA and SCA (Table II). These
results proposed that both additive and non-additive gene
actions were complicated in controlling pod length. Parent
G6 acquired maximum GCA (0.89) and was considered the
best general combiner for pod length, as indicated in the
Table III.
Among the F2 populations G4 G6 was the best specific
combiner, as it indicated the maximum SCA (0.93)
followed by G2 G6 (0.57), as showed in the Table IV. The
maternal effects were also highly significant for pod length.
Cross combination G9 G6 obtained the desire maximum
RCA (0.67) followed by G4 G2 (0.33), as indicated in the
Table V. The maximum reciprocal effects showed by these
cross combinations for pod length were because of
cytoplasmic genes donated by the female parents. Our
results are in agreement to the earlier findings of Rameeh
(2010) in Brassica napus L. who found significant GCA
and SCA effects for pod length and revealed the importance
of both types of gene actions. Similarly, Maurya et al.
(2012) found significant GCA effects in Brassica juncea
showed the presence of additive gene action for pod length.
Sabaghnia et al. (2010) reported significant SCA effects for
pod length in rapeseed and Arifullah et al. (2011) reported
significant SCA effects in Brassica juncea L. indicated the
presence of non-additive gene action for pod length.
Genetic and environmental variances for pod length were
1.21 and 0.44, respectively. The estimated heritability (bs)
was high i.e. 0.73 (Table VI). Chaghakaboodi et al. (2012)
also reported high heritability in Brassica napus L.
However, Aytac and Kinaci (2009) and Zare and
Sharafzadeh (2012) found low broad sense heritability for
pod length in rapeseed (Brassica napus L.).

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10

Table II
Mean squares for ANOVA and combining ability for various important traits in a 44 diallel cross of Brassica napus conducted at The University of Agriculture,
Peshawar during 2012-13

ANOVA

Parameters
GMS
Days to 50% flowering
Plant height
Main raceme length
Pod length

EMS

COMBINING ABILITY
CV%

GCA
2.95

SCA

RCA

ns

12.46*

8.63ns

27.07*

13.08

4.06

346.56**

53.5

5.32

171.20**

165.85**

37.35ns

87.96**

20.91

8.92

60.49**

28.93**

14.12 ns

4.08**

0.44

11.88

2.90**

1.11**

0.71**

** Significant at P0.01, * significant at P0.05 and ns = Non significant

Table III
Estimates of GCA effects for various important traits in a 44 diallel cross of Brassica napus conducted at The University of Agriculture, Peshawar during 201213

Parameter
Days to 50% flowering
Plant height
Main raceme length
Pod length

G2

G4

G6

G9

-0.78
-4.98
-1.28
-0.41

0.33
-1.92
-3.25
-0.33

-0.15
5.89
1.95
0.89

0.6
1.0
2.58
-0.15

Table IV
Estimates of SCA effects for various important traits in a 44 diallel cross of Brassica napus conducted at The University of Agriculture, Peshawar during 201213

Parameters
Days to 50% flowering
Plant height
Main raceme length
Pod length

G2G4

G2G6

G2G9

G4G6

G4G9

G6G9

3.24
9.76
-0.08
-0.53

-1.58
5.13
3.9
0.57

2.7
-1.51
-3.3
-0.51

-0.27
5.39
2.25
0.93

-2.17
4.65
-3.4
0.13

-2.17
4.65
-3.4
0.13

Table V
Estimates of RCA effects for various important traits in a 44 diallel cross of Brassica napus conducted at The University of Agriculture, Peshawar during 201213

Parameters

G4G2

G6G2

G6G4

G9G2

G9G4

G9G6

-0.53

3.38

2.05

-0.1

3.1

-0.62

5.4

0.52

5.4

-0.88

1.14

7.17

Main raceme length

2.1

-1.39

5.3

-1.88

-1.92

0.85

Pod length

0.33

-0.98

0.13

0.15

-0.76

0.67

Days to 50% flowering

Plant height

Table VI
Variance components and heritability(BS) for various important traits in a 44 diallel cross of Brassica napus conducted at The University of Agriculture, Peshawar
during 2012-13

Parameters
Days to 50% flowering
Plant height
Main raceme length
Pod length

Vg
4.66
97.68
22.35
1.21

Ve
13.08
53.5
20.91
0.44

Vp
17.74
151.18
43.26
1.65

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h2 (bs)
0.26
0.65
0.52
0.73

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CONCLUSIONS AND RECOMMENDATIONS
From the present research work to estimate combining
ability and heritability in Brassica napus L. genotypes
the following conclusions have been derived;

[14]

[15]

Analysis of variance showed significant differences for

the genotypes. Parental genotypes G6 and G9 were best
general combiners whereas cross combinations G2 G4
and G6 G4 were best specific and reciprocal
combinations, respectively therefore could be used in
future breeding program. Plant height and main raceme
length were high heritable traits and have the room for
further improvement in the future breeding programs. The
overall study discloses the prominence of both additive
and non-additive genetic variability suggesting the use of
integrated breeding strategies which can efficiently
exploit the additive as well as non-additive genetic
effects.

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[17]

[18]

[19]

[20]

REFERENCES
[1]

[16]

Akbar, M., B. Tahira, B.M. Atta and M. Hussain. 2008.

Combining ability studies in rapeseed (Brassica napus L.).
Int. J. agric. & biol. 10(2): 205-208.
Ali, N., F. Javidfar and A.A. Attary. 2002. Genetic
variability, correlation and path analysis of yield and its
components in winter rapeseed (Brassica napus L.). Pak. J.
Bot. 34(2): 145-150.
Ali, N., F. Javidfar, J.Y. Elmira and M.Y. Mirza. 2003.
Relationship among yield components and selection criteria
for yield improvement in winter rapeseed (Brassica napus
L.). Pak. J. Bot. 35(2): 167-174.
Ali. Y., Farhatullah, H. Rahman, A. Nasim, S.M. Azam and
A. Khan. 2013. Heritability and correlation analysis for
morphological and biochemical traits in Brassica carinata.
Sarhad J. Agric. 29(3): 359-370.
Aslam K.S., H. Ahmad, A. Khan, M. Saeed, S.M. Khan and
B. Ahmad. (2009). Using line tester analysis for earliness
and plant height traits in Sunflower (Helianthus Annus L.).
Recent Research in Science and Technology, 1(5): 202206.
Oghan, A.H., M.H. Fotokianb, F. Javidfar and B.
Alizadeha.2009. Genetic analysis of seed
yield, days to
flowering and maturity in oilseed rape (Brassica napus L.)
using diallel crosses. Int. J. Plant Prod. 3(2):19-26.
Arifullah. M., M. Munir, A. Mahmood, S.K. Ajmal and G.
Shabbir. 2012. Combining ability analysis of some yield
attributes in Indian mustard (Brassica juncea L.). Pak. J.
Agric. Res. 25(2): 104-109.
Aytac, Z. and G. Kinaci. 2009. Genetic variability and
association studies of some quantitative characters in winter
rapeseed (Brassica napus L.) Afric. J. Biotechnol. 8(15):
3547-3554.
Azizinia, S. 2011. Combining ability analysis for yield
component parameters in winter rapeseed genotypes
(Brassica napus L.). J. Oilseed Brassica, 2(2): 67-75.
Chaghakaboodi, Z., D. Kahrizi and A. Zebarjadi. 2012.
Heritability and genetic advance in rapeseed (Brassica
napus L.). Iran. J. Genet. & Plant Breed. 1(2): 16-21.
Dar, Z.A., S.A, Wani, G.M. Habib, G. Ali, P.A. Sofi, S.A.
Dar and A.M. Iqbal. 2013. Analysis of combining ability in
brown sarson (Brassica rapa L.) under temperate
conditions. Afric. J. Agric. Res. 8(117): 1603-1607.
Griffing, B. 1956. Concept of general and specific
combining ability in relation to diallel systems. Aust. J.
Biol. Sci. 9: 463-493.
Gupta, P., Chaudhary and S.K. Lal. 2011. Heterosis and
combining ability analysis for yield and its components in

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

11

Indian mustard (Brassica juncea L. Czern & Coss).

Acadmic J. Plant Sci. 4(2): 45-52.
Huang, Z., P. Laosuwan, T. Machikowa and Z. Chen. 2009.
Combining ability for seed yield and other characters in
rapeseed. Suranaree J. Sci. Tech. 17(1): 39-47.
Mahak, S., A. Tomar, C.N. Mishra and S.B.L. Srivastava.
2011. Genetic parameters and character association studies
in Indian mustard. J. Oilseed Brassica, 2: 35-38.
Marjanovic, J. A., R. Marinkovic, S. Ivanovska, M.
Jankulovska, A. Mijic and N. Hristov. 2011. Variability of
yield determining components in winter rapeseed (Brassica
napus L.) and their correlation with seed yield. Genetika,
43(1): 51-66.
Maurya, N., A.K. Singh and S.K. Singh. 2012. Analysis of
combining ability in indian mustard (Brassica juncea L.).
Indian J. Plant Sci. 1(2-3): 116-123.
Nasim, A. Farhatullah, S. Iqbal, S. Shah and S.M. Azam.
2013. Genetic variability and correlation studies for morphophysiological traits in Brassica napus L. Pak. J. Bot. 45(4):
1229-1234.
PBS, 2012. Pakistan Bureau of Statistics (2011-2012).
Pakistan Oilseed Development Board. Government of
Pakistan, 22-23.
Panhawar, S.A., M.J. Baloch,, W.A. Jatoi, N.F. Veesar and
M.S. Majeedano. (2008). Combining ability estimates from
line tester mating design in upland cotton. Proceeding.
Pakistan Academy of Science, 45(2): 69-74.
Patel, P.J. and S.R. Vyas. 2011. Heritability and genetic
advance for yield and quality traits in Indian mustard [
Brassica juncea (L.) Czern and Coss]. Adv. Res. J. Crop
Improv. 2(2): 212-214.
Rameeh V. 2012. Combining ability and heritability
estimates of main agronomic characters in rapeseed
breeding lines using line tester analysis. J. Agric. Sci.
57(3): 111-120.
Rameeh, V. 2011. Combining ability analysis of rapeseed
genotypes under restricted nitrogen application. J. Oilseed
Brassica, 2(1): 7-12.
Rameeh, V. 2010. Combining ability and factor analysis in
F2 diallel crosses of rapeseed varieties. Plant Breed. Seed
Sci. 62(1): 73-83.
Sabaghnia, N., H. Dehghani, B. Alizadeh and M.
Mohghaddam. 2010. Heterosis and combining ability
analysis for oil yield and its components in rapeseed. Aust.
J. Crop Sci. 4(6): 390-397.
Shehzad, F.D. and Farhatullah. 2012. Selection of elite
genotypes for yield and associated traits in in F2-3 families
of interspecific crosses in brassica species. Pak. J. Bot.
44(4): 1297-1301.
Singh, M., A. Tomar, C.N. Mishra and S.B.L. Srivastava.
2011. Studies on genetic components for seed yield and its
contributing traits in Indian mustard (Brassica juncea). J.
Oilseed Brassica, 2(2): 83-86.
Singh R.K. and B.D. Chowdhury. (1985). Biometrical
method in quantitative genetic analysis 2nd Edn, Kalyani
Publishers, Ludhiana, New Dehli, India, pp: 54-57.
Singh, M., L. Singh and S.B.L. Srivastava. 2010.
Combining ability analysis in Indian mustard (Brassica
juncea L. Czern & Coss). J. Oilseed Brassica, 1(1): 23-27.
Sprague, G.F. and L.A. Tatum. 1942. General versus
specific combining ability in single crosses of corn. J.
Amer. Soc. Agron. 34: 923-32.
Zare, M. and S. Sharafzadeh. 2012. Genetic variability of
some rapeseed (Brassica napus L.) cultivars in Southern
Iran. Afric. J. Agric. Res. 7(2): 224-229.

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