Eco. Env. & Cons. 22 (1) : 2016; pp.

(345-349)
Copyright@ EM International
ISSN 0971–765X

Combining ability analysis and gene action for yield

and its contributing traits in tomato (Solanum
lycopersicum L.) under North Western
Himalayan region
Nidhish Gautam, 1Manish Kumar, 1Sandeep Kumar, 1Amit Vikram, 2RK Dogra and 3N Bharat
1

1
Department of Vegetable Science,
2
Department of Fruit Science,
3
Department of Seed Science and Technology,
Dr YS Parmar University of Horticulture and Forestry, Nauni, Solan 173 230 HP, India

(Received 6 July, 2015; accepted 30 August, 2015)

ABSTRACT
Diallel analysis revealed highly significant differences among tomato genotypes for days to maturity, plant
height, number of fruits per plant, fruit weight, and fruit yield per plant. Significant mean squares for
general combining ability (GCA) and specific combining ability (SCA) indicated joint role of additive, non-
additive for the expression of days to maturity and fruit yield per plant. The predictability ratio of GCA/
SCA variance was found less than one for days to maturity, plant height, number of fruits per plant, fruit
weight, and fruit yield per plant showing preponderance of non-additive gene effects. Among parents,
UHFT-10 and Solan Lalima were found good general combiner for yield and some of the yield related traits
studied. The hybrids viz. UHFT-55 x Solan Lalima, UHFT-9 x Solan Lalima and UHFT-10 x Solan Lalima
had significant SCA effects for yield and were suggested for the exploitation of heterosis.

Key words : Diallel (excluding reciprocals), Analysis of variance (ANOVA), Gene action, General Combining Ability (GCA),
Specific combining ability (SCA).

Introduction annual production of 18.22 million tonnes (Anony-

Tomato (Solanum lycopersicum L.) belongs to the
family Solanaceae. It is grown as summer as well as
off season vegetable in Himachal Pradesh. Tomato
is a rich source of vitamin A, C and minerals like Ca,
P and Fe (Dhaliwal et al., 2003). Tomatoes are major
contributors of antioxidants such as carotenoids (es-
pecially, lycopene and β-carotene), phenolics, ascor-
bic acid (vitamin C) and small amounts of vitamin E
in daily diets (Rai et al., 2012). In India, tomato is
grown on an area of 0.879 million hectares with an
mous, 2013). In Himachal Pradesh, total area under
tomato is 993 hectares with the production of 413710
tones (Anonymous, 2013). Current open pollinated
(OP) varieties of tomato are unable to meet the do-
mestic demand due to their low genetic potential,
susceptibility to biotic and abiotic stresses, limited
area under cultivation, water shortage and competi-
tion with major crops (Saleem et al., 2011; Sajjad et al.,
2011; Akhtar et al., 2012). Hybrid variety (F1 popula-
tion) gives 3 to 4 times more yield in contrast to that
of OP variety (Tiwari and Choudhury, 1986). Unlike

*Corresponding author’s email : nidhish635@gmail.com

346
OP varieties, the superior characters of F1 hybrids
however, are lost during further generations there-
fore; growers need to buy single generation hybrid
seed every time when they want to plant. Main rea-
son for slow progress in tomato hybrid breeding in
India is lack of good combiner parents to be crossed
for exploitation of heterosis. India is facing higher
imports of tomato seed due to limited quality seed
producing agencies that can fulfill domestic seed re-
quirements.
Different biometrical techniques are now avail-
able to select successful parent lines suitable for hy-
brid seed production. Diallel analysis technique de-
veloped and illustrated by Hayman (1957) and Jinks
(1956) provides guideline for the assessment of rela-
tive breeding potential of the parents and has been
extensively used to identify good combiner parents
in various crops like hot pepper (Legesse, 2001), to-
mato (Chishti et al., 2008) and okra (Wammanda et
al., 2010). This technique provides information on
gene action controlling the expression of desired
traits. Based on this information on combining abil-
ity and gene action, the desirable selected lines can
be combined either to exploit hybrid vigor by accu-
mulating non-additive gene effects or to evolve cul-
tivars by accumulating additive gene effects. The
rationale of the present study was to pick elite lines
of tomato to develop hybrids suitable for field culti-
vation using diallel analysis (Griffing, 1956). Having
recognition of such lines, hybrid varieties of tomato
can be produced on commercial scale to increase
yield, supply quality seed to farmers at low cost.
Materials and Methods
Five tomato lines viz., UHFT-22, UHFT-55, UHFT-9,
UHFT-10, EC-2798 and Solan Lalima were crossed
in diallel fashion (excluding reciprocals) following
Table 1. Analysis of variance for combining ability for various traits in tomato
Source > Mean Sum of Squares
Trait GCA
df (Degree of freedom) 15
>
Days to first flowering 63.71*
Days to marketable maturity 90.86*
Plant height (cm) 13012.76*
Average fruit weight (g) 1108.03*
Number of fruits per plant 28.61*
Yield per plant (kg) 1.56*
Eco. Env. & Cons. 22 (1) : 2016
Griffing (1956). The 21 genotypes (15 direct F 1
crosses + 6 parents) were grown in Experimental
Research Farm, Department of Vegetable Science,
Dr Y S Parmar, UHF, Nauni, Solan (HP), India fol-
lowing randomized complete block design with 3
replications during 2012-13. Twenty eight days old
nursery seedlings were transplanted at a distance of
90 cm × 30 cm. Twelve plants of each genotype per
replication were grown by adopting standard agro-
nomic practices to maintain healthy crop stand. The
data were recorded on days to maturity, plant
height (cm), number of fruits per plant, fruit weight
(g), and fruit yield per plant (kg). Analysis of vari-
ance was performed according to Steel et al. (1997).
Combining ability (general combining ability re-
ferred as GCA, specific combining ability referred as
SCA) analysis was carried out following Model-²,
Method-²² of Griffing (1956).
Results and Discussion
Analysis of variance indicated significant differ-
ences among all the genotypes for days to maturity,
plant height, number of fruits per plant, and fruit
yield per plant as reported earlier (Saleem et al.,
2013). The analysis of variance for combining ability
divided genetic variation into GCA and SCA com-
ponents. Mean sum of squares from the analysis of
GCA and SCA were found significant for traits un-
der study (Table 1). The proportions of GCA and
SCA were estimated to assess the relative impor-
tance of GCA and SCA in the expression of different
traits (Table 2). The Magnitude of SCA variance was
greater than that of GCA variance for traits days to
maturity, plant height, number of fruits per plant,
fruit weight and fruit yield per plant indicating the
major control of non-additive type of gene action. In
tomato, Saidi et al., (2008) reported the importance
SCA Errors Total
5 40 60
11.98* 0.78 76.47
18.43* 1.17 110.46
1728.21* 10.66 14751.63
147.27* 2.13 1257.43
18.77* 1.00 48.38
0.33* 0.01 1.90
GAUTAM ET AL 347

of additive effects for plant height, additive and
non-additive effects for number of fruits per plant
and fruit weight, while dominance effects for fruit
yield per plant. However, Saleem et al., (2009) re-
ported non-additive effects for fruit weight, fruit
length, fruit width, number of fruits per plant and
fruit yield per plant.
In perusal to GCA effects of the parent lines
(Table 3), UHFT-22 showed desirable GCA effects
for days to marketable maturity, days to first flow-
ering and plant height as it had GCA values of -3.43,
-2.90 and -46.73, respectively. Lines, UHFT-9 and
Solan Lalima had higher GCA effect for average
fruit weight with GCA value of 7.53 and 5.89 respec-
tively. Lines appeared better for number of fruits
per plant (UHFT-10) and plant height (Solan Lalima)
with GCA value of 1.12 and 7.23, respectively. Sig-
nificant positive GCA effect for fruit yield per plant
were exhibited by Solan Lalima and UHFT-10 as it
had GCA values of 0.23 and 0.23, respectively. Simi-
lar results had already been reported elsewhere by
various researchers in tomato (Ahmad et al., 2009
and Singh et al., 2010). General combining ability has
direct relationship with narrow sense heritability
and represents fixable portion (additive and addi-
tive x additive interaction) of genetic variation, thus
helps in selection of parents suitable for hybridiza-
tion (Geleta et al., 2006 and Saleem et al., 2009) to
develop cultivars with desired traits of interest.
Lines Solan Lalima, UHFT-9 and UHFT-10 were
rated as best general combiner and can be used as
donors for yield and some other traits to develop
early maturing and high yielding genotypes
through multiple crossing programme.
Specific combing ability effects of hybrids have
been presented in table 4. Certain hybrids viz.,
UHFT-55 × EC 2798 had desirable SCA value of -
4.89 followed by UHFT-22 × EC 2798 (-3.38), and
UHFT-22 × Solan Lalima (-2.23) for the development
of early maturing hybrids. Among all the cross com-
binations, hybrids namely UHFT-55 × Solan Lalima
had highest positive SCA effect for plant height
(52.76) followed by UHFT-55 x UHFT-22 (52.66),
EC-2798 × Solan Lalima (22.73), UHFT-10 × Solan
Lalima (19.68), UHFT-9 × UHFT-10 (18.19), and
UHFT-9 × Solan Lalima (9.92). Positive SCA effects
for high fruit weight were exhibited by hybrid
UHFT-9 × EC 2798 with SCA value of 8.04, UHFT-

Table 2. Estimates of genetic components of variance for various traits in tomato

Trait σ2 GCA σ2 SCA σ2 g σ2 s σ2g/σ2s
(Variance Ratio)
Days to first flowering 21.24 3.99 2.56 3.22 0.80
Days to marketable maturity 30.29 6.14 3.64 4.97 0.73
Plant height (cm) 4337.59 576.07 540.87 565.41 0.96
Average fruit weight (g) 369.35 49.09 45.90 46.96 0.98
Number of fruits per plant 9.54 6.26 1.07 5.25 0.20
Yield per plant (kg) 0.52 0.11 0.06 0.10 0.64

Table 3. Estimates of general combining ability effects of parents for various traits in tomato
CharacterParents Days Days to Plant Average Number Yield
to first marketable height fruit of fruits per plant
flowering maturity (cm) weight (g) per plant (kg)
UHFT-55 -0.60* -0.78* 11.66* -1.79* 0.49* -0.03
UHFT-22 -2.90* -3.43* -46.73* -10.68* -1.72* -0.42*
UHFT-9 1.57* 1.90* 7.98* 7.53* 0.18 0.23*
UHFT-10 0.86* 0.96* 8.47* 2.84* 1.12* 0.15*
EC-2798 1.13* 1.41* 2.90* -3.78* -0.89* -0.16*
Solan Lalima -0.06 -0.06 15.73* 5.89* 0.82* 0.23*
SE (gi) 0.16 0.20 0.61 0.27 0.19 0.02
SE (gi-gj) 0.26 0.32 0.94 0.42 0.29 0.03
CD (gi) 0.33 0.42 1.27 0.56 0.40 0.04
CD (gi-gj) 0.54 0.67 1.96 0.87 0.60 0.06
*Significant at 5% level of significance
348 Eco. Env. & Cons. 22 (1) : 2016

10 × EC 2798 (7.26), UHFT-9 × Solan Lalima (5.08),
UHFT-22 × UHFT-10 (4.73), and UHFT-55 × Solan
Lalima (4.66). Five hybrids viz., UHFT-55 × Solan
Lalima, EC 2798 × Solan Lalima, UHFT-9 × Solan
Lalima, UHFT-22 × EC 2798, and UHFT-10 × Solan
Lalima had desirable SCA effect of 3.05, 3.01, 2.17,
1.74, and 1.59, respectively for higher number of
fruits per plant. Six hybrids viz., UHFT-55 × Solan
Lalima (0.36) followed by UHFT-9 × Solan Lalima
(0.34), UHFT-10 × EC 2798 (0.28), UHFT-9 × EC 2798
(0.28), UHFT-55 × UHFT-22 (0.19) and UHFT-55 ×
EC 2798 (0.19) had desirable SCA effects for yield
per plant.
According to Singh and Narayanan (2004), a SCA
effect refers to non-additive gene action (mainly
dominance, interactions of dominance x dominance,
additive x dominance and non-allelic loci) and has
positive relationship with heterosis. Therefore, hy-
brid UHFT-9 x Solan Lalima, UHFT-10 x Solan
Lalima and UHFT-55 x Solan Lalima were rated as
the best crosses for improvement in yield and some
of its contributing traits. Heterosis breeding was rec-
ommended for those combinations, which had sig-
nificantly high SCA effects for fruit yield per plant.
Normally SCA effect does not contribute to the im-
provement of self-pollinated crops like tomato.
However, the cross showing significant SCA value,
provided that at least one of its parents had high
desirable GCA effect, is expected to produce favor-
able transgrassive segregants through hybridization
with delayed selection (because of dominance and
epistatic interactions) if the complementary epistatic
effect present in that cross acts in the same direction
to maximize desirable plant attributes (Salimath &
Bahi, 1985).
Conclusion
From the present investigations, it can be concluded
that tomato lines UHFT-9 and Solan Lalima proved
as best general combiner and could be utilized in
multiple crossing program to develop early matur-
ing and high yielding tomato genotypes. Three
crosses viz., UHFT-9 x Solan Lalima, UHFT-10 x
Solan Lalima and UHFT-55 x Solan Lalima had at
least one good general combiner parent and had
high SCA effect, thus suggesting heterosis breeding
as a valid strategy for the development of vigorous

Table 4. Estimates of specific combining ability effects of crosses for various traits in tomato
CharacterParents Days to first Days to Plant Average Number Yield
flowering marketable height fruit weight of fruits per plant
maturity (cm) (g) per plant (kg)
UHFT-55 X UHFT-22 1.28* 1.62* 52.66* 4.33* 1.39* 0.19*
UHFT-55 X UHFT-9 0.77 1.05 -7.94* -7.98* 0.09 -0.23*
UHFT-55 X UHFT-10 -1.55* -2.67* -6.18* 0.21 0.92 0.07
UHFT-55 X EC-2798 -4.12* -4.89* -5.89* 3.89* 1.47* 0.18*
UHFT-55 X Solan Lalima -0.70 -0.72 -15.80* 4.66* 3.05* 0.36*
UHFT-22 X UHFT-9 -0.42 -0.53 -29.57* 2.67* 1.27* 0.14*
UHFT-22 X UHFT-10 -1.25* -1.42* -32.10* 4.73* 0.26 0.13*
UHFT-22 X EC-2798 -2.78* -3.38* -23.52* 2.94* 1.74* 0.16*
UHFT-22 X Solan Lalima -1.86* -2.23* -25.78* -12.89* -3.07* -0.55*
UHFT-9 X UHFT-10 0.31 0.48 18.19* 4.46* 0.03 0.16*
UHFT-9 X EC-2798 0.64 0.76 -2.76 8.04* 0.51 0.25*
UHFT-9 X Solan Lalima -1.30* -1.83* 9.92* 5.08* 2.17* 0.34*
UHFT-10 X EC-2798 -0.45 -0.47 -1.50 7.26* 1.17* 0.28*
UHFT-10 X Solan Lalima 0.88 1.17* 19.68* 1.26 1.59* 0.16*
EC-2798 X Solan Lalima 1.54* 1.82* 22.73* -0.02 3.01* 0.16*
SE (sij) 0.45 0.55 1.67 0.75 0.51 0.05
SE (sij-sik) 0.88 1.09 3.25 1.47 1.01 0.10
SE (sij-skl) 0.62 0.77 2.31 1.03 0.71 0.07
CD (sij) 0.94 1.14 3.47 1.56 1.06 0.10
CD (sij-sik) 1.83 2.27 6.76 3.06 2.10 0.21
CD (sij-skl) 1.29 1.60 4.80 2.14 1.48 0.15
*Significant at 5% level of significance
GAUTAM ET AL
high yielding genotypes from the succeeding prog-
enies.
References
Ahmad, S., Quamruzzaman, A.K.M. and Nazim-Uddin,
M. 2009. Combining ability estimates of tomato
(Solanum lycopersicum) in late summer. SAARC J.
Agri. 7 (1) : 43-56.
Akhtar, K.P., Saleem, M.Y., Asghar, M., Ali, S., Sawar, N.
and Elahi, M.T. 2012. Resistance of solanum species
to phytophthora infestans evaluated in the de-
tached-leaf and whole-plant assays. Pak. J. Bot. 44 (3)
: 1141-1146.
Anonymous. 2013. www.nhb.co.in (National Horticulture
Board Database)
Chishti, S.A.S., Khan, A.A., Sadia, B. and Khan, I.A. 2008.
Analysis of combining ability for yield, yield com-
ponents and quality characters in Tomato. J. Agri.
Res. 46 (4) : 325-331.
Dhaliwal, M.S., Singh, S. and Cheema, D.S. 2003. Line x
tester analysis for yield and processing attributes in
tomato. J. Res. 40 (1): 49-53.
Geleta, L.F. and Labuschagne, M.T. 2006. Combining abil-
ity and heritability for vitamin C and total soluble
solids in pepper (Capsicum annuum L.). J. Sci. Food.
Agric. 86 : 1317-1320.
Griffing, B. 1956. Concept of general and specific combin-
ing ability in relation to diallel crossing system. Aust.
J. Biol. Sci. 90 : 463-492.
Hayman, B.I. 1957. Interaction, heterosis and diallel
crosses. Genet. 42 : 336-355.
Inamullah., Mohammad, F., Din, S. and Gull, R. 2006.
Diallel analysis of the inheritance pattern of agro-
nomic traits of bread wheat. Pak. J. Bot. 38(4) : 1169-
1175.
Jinks, J.L. 1956. The F2 and backcross generations from a
set of diallel crosses. Heredity. 10 : 1-30.
Legesse, G. 2001. Combining ability study for green fruit
yield and its components in hot pepper (Capsicum
annuum L.). Acta Agron. Hung. 48 (4) : 373-380.
Rai, G.K., Kumar, R., Singh, A.K., Rai, P.K., Rai, M.,
Chaturvedi, A.K. and Rai, A.B. 2012. Changes in
antioxidant and phytochemical properties of tomato
349
(Lycopersicon esculentum mill.) under ambient condi-
tion. Pak. J. Bot. 44 (2): 667-670.
Saidi, M., Warade, S.D. and Prabu, T. 2008. Combining
ability estimates for yield and its contributing traits
in tomato (Lycopersicon esculentum). Int. J. Agri. Bio.
10: 238-240.
Sajjad, M., Ashfaq, M., Suhail, A. and Akhtar, S. 2011.
Screening of tomato genotypes for resistance to to-
mato fruit borer (Helicoverpa armiger, Hubner) in
Pakistan. Pak. J. Agri. Sci. 48 : 59-62.
Saleem, M.Y., Akhtar, K.P., Asghar, M., Iqbal, Q. and
Rehman, A. 2011. Genetic control of late blight, yield
and some yield related traits in tomato (Solanum
lycopersicum L.). Pak. J. Bot. 43 (5): 2601-2605.
Saleem, M.Y., Asghar, M. and Iqbal, Q. 2013. Augmented
analysis for yield and some yield components in
tomato (Lycopersicon esculentum Mill.). Pak. J. Bot. 45
(1) : 215-218.
Saleem, M.Y., Asghar, M., Haq, M.A., Rafique, T., Kamran,
T. and Khan, A.A. 2009. Genetic analysis to identify
suitable parents for hybrid seed production in to-
mato (Lycopersicon esculentum Mill.). Pak. J. Bot.
41 (3) : 1107-1116.
Salimath, P.M. and Bahl, P.N. 1985. Heterosis and combin-
ing ability for earliness in chickpea (Cicer arietinum
L.). Indian J. Genet. 45 : 97-100.
Singh, P. and Narayanan, S.S. 2004. Biometrical Tech-
niques in Plant Breeding. Kalyani Publ., New Delhi,
India.
Singh, S.P., Thakur, M.C. and Pathania, N.K. 2010. Recip-
rocal cross differences and combining ability stud-
ies for some quantitative traits in tomato
(Lycopersicon esculentum Mill.) under mid hill condi-
tions of western Himalayas. The Asian J. Hort. 4 (2):
473-477.
Steel, R.G.D., Torrie, J.H. and Dick, D.A. 1997. Principles
and Procedures of Statistics-a Biometrical Approach.
McGraw Hill Book Co., New York.
Tiwari, R.N. and Choudhury, B. 1986. Tomato. In: Solana-
ceous Crops. (Eds.): B. Som & K.N. Prokash. Calcutta,
pp. 224-280.
Wammanda, DT., Kadams, A.M. and Jonah, P.M. 2010.
Combining ability analysis and heterosis in a dial-
lel cross of okra (Abelmoschus esculentus L. Moench).
African J. Agric. Res. 5(16) : 2108-2115.