Vegetable Science (2014) 41 (1) : 37-41
Genetic analysis of yield and yield attributing traits in bitter gourd
Anjali Shukla, Umesh Singh, A.K. Rai, D.R. Bharadwaj and Major Singh
Received : April 2014 / Accepted : June 2014
Abstract
Gynoecious bitter gourd has the potential for increasing
yield in commercial cucurbitaceous. Therefore, a generation
means analysis was conducted to investigate the inheritance
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of yield attributing traits in crosses of gynoecious
monoecious. Progeny (F1, F2, and F3) from a cross between
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gynoecious lines (Gy333 and Gy323) and monoecious line
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DRAR-1were used in the present study. The additive
dominance and dominance dominance interaction effects
were noted in most of the crosses for the entire traits.The
joint scaling test showed that cross Gy333 DRAR-1 and
Gy323DRAR-1 showed significant epistatic gene effects.
2 value was significant for all traits except days to first
flowers and anthesis, plant height, internodal length, fruit/
plant seeds/fruit and yield/plant showed non-significant
2value hence the additive dominance model.In cross
Gy323 DRAR-1, 2 value was significant for all traits except
yield/ plant. Both additive and dominance components were
significant for most of traits.These traits showed duplicate
type of epistasis for the above crosses.The development of
early flowering genotypes possessing desirable fruit weight
characteristics, however, will likely be complicated by
inheritance of gynoecious in dominance and epistatic
effects. Positive additive additive is indicating the
possibility of obtaining transgressive segregants in later
generations, whereas dominance dominance gene effects
indicated a duplicate epistasis which will be undesirable for
selection and genetic improvement in gynoecious bitter
gourd.
Keywords: bitter go urd, gynoecious, qu anti tati ve
inheritance, gene action, epistasis.
Anjali Shukla, Umesh Singh, D.R. Bharadwaj and Major Singh
Division of Crop Improvement, Indian Institute of Vegetable
Research, PB.- 01, Jakhini (Shahanshapur), Varanasi, Uttar
Pradesh-221 305, India
A.K. Rai
Professor, Centre of Advanced Studies, Department of Botany,
Banaras Hindu University, Varanasi, Uttar Pradesh 221 005,
India
Introduction
Bitter gourd (MomordicacharantiaL., 2n= 22) is an
economically important, allogamy, vegetable species,
which is used in traditional or folk medicine to treat
diabetes. The bitterness of most cucurbits is mainly due
to cucurbitacins. The bitterness of bitter gourd is due to
the cucurbitacin-like alkaloid momordicine and triterpene
glycosides (momordicoside K and L) (Behraet al.,
2010).Inheritance of gynoecism in bitter gourd was
studied in a 100% gynoecious line (Gy323 and Gy333).
The F2 and testcross segregation data revealed that
gynoecism in Gy323 and Gy333 is under the control of
a single, recessive gene (Ram et al., 2006).
High yield and uniform fruit shape, size and excellent
quality are prerequisites for the release of superior melon
varieties. Gynoecious for yield and/or its associated
components has been reported in melon (Abdalla and
Aboul-Nasr,2002).Lippert and Legg (1972) evaluated the
gene action of yield traits in melon, and determined that
additive and non-additive variance components were
important in the genetic control of yield-associated traits.
However, the relative importance of additive, dominant
and epistatic contributions was not reported, and other
studies that evaluate gene action controlling such traits
in bitter melon do not exist.
Generation means analysis (GMA; Mather and Jinks,
1982) has been used successfully to study the genetics
of melon in term of gynoecious. The generation mean
analysis is commonly employed in studies of inheritance
of quantitative traits. Given the lack of genetic information
related to yield components in gynoecious for bitter
melon, a GMA study was designed todetermine gene
action, estimate components of variance and calculate
the minimum number of effective factors of several yield
components in gynoecious bitter melon. In relation to
direction of dominance, reciprocal effects and epistasis,
relevant differences between the polygenic systems were
evidenced. An assessment of these genetic parameters
 38 Shukla et al. : Genetic analysis of yield and yield attributing traits in bitter gourd
will allow for the development of efficient breeding
strategies and new hybrid (Gynoecious monoecious)
for gynoecious bitter melon cultivar improvement.
Materials and Methods
Experimental parents were selected based on their higher
yield, earliness and yield attributing traits. Gynoceious
and monoceious bitter gourd parents used were highly
inbred lines. Gy323 and Gy333 are gynoecious, in which
Gy323 with Large vines, large size leaves with heart
shape, while, Gy333 gynoecious plant, medium vines,
more branching, profuse bearing, green fruist both
developed by IIVR (Ram et al., 2002). The fractal
architecture of DRAR-1 is monoecious, large vines,
medium leaves, less branching, light green fruit
developed by TNAUfor five generations before use.
C ross es were ma de betwe en gynoc eious and
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monoceious parents as follows: Gy323 DRAR-1 and
Gy333 DRAR-1. Generation means analysis was
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performed using each gynoceious (P1), monoceious
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parent (P2), F1, F2 and F3 generations. All crosses were
controlled pollinations and experiment with 3 different
parents and their generationwere conducted at the
Indian Institute of Vegetable Research, Varanasi, India.
The crop was planted in rows 2.5 mapart with spacing
of 45 cm between plants. All the recommended
agronomical practices along with plant protection
measures were followed to grown a successful crop.
The population segregating was characterized for eleven
traits viz.,1st flower anthesis, plant height (cm),
internodal length (cm), number of branches/plant, days
to first picking, fruit weight (g), fruit length (cm), fruit
diameter (cm), fruits/plant, seeds/fruit, yield/plant (g)
were evaluated based on evaluation system. In each
replication ten plants in each population were marked
for observations. Within each block the segregating
generations were replicated three and five times,
respectively.
Analyses of generations by GMA were conducted using
an additivedominance model (Cavalli, 1952), non-
weighted scaling test method based on a five-parameter
model and using sequential parameter model fitting
(Kearsey and Pooni, 1996). The average gene effects
expectations of the five basic generations are given as
proposed by Mather and Jinks (1982).5-parameter
models and were used to estimate the gene effects (m,
d & h) and their interaction (l &i) following Hayman
(1858). Thestandard error of each parameter was
calculated as proposed by Lynchand Walsh (1998).
Result and Discussion
The development of gynoecious bitter gourd cultivars
having potential to produce optimum yields is highly
desirable where monoecious bitter gourd is becoming
very limited yield. To detect the relative importance of
components of genetic variation, additive (d), dominance
(h) and contribution (I &l) were estimated by partitioning
the population means of parents, F 1, F 2 and F 3.
Relativegynoceious magnitude of additive component
D was significant for most of the characters except
branches/plant indicated that the additive components
of genes played an important role in gynoecious related
characters expression and on their inheritance.
In the present study, there are two crosses Gy333
DRAR-1 and Gy323 DRAR-1 used in generation.5
generation (P1, P2, F1, F2& F3) of 2 cross combinations
have been analyzed for 11 characters and the gene
effects we re e stimated through pa rtitioning the
generation means into different components. In the event
of inadequacy of the model the data was refitted for 3
parameter models. For the characters where all the 3
and 5 components were not significant in the both
paramete r model the elimination of the gene tic
component was also done to increase the precision of
the results. The negative estimates of dominance gene
effects observed for flower anthesis implied that the
inheritance of these characters of gynoecioustend
toward the lower parent. Additive Additive interaction
estimates were mostly negative for these characters
effectively.
Cross Gy333 DRAR-1 showed significant epistatic
gene effects (Table 1). 2 value was significant for all
traits except days to first flowers and anthesis, plant
height, internodal length, fruit/plant seeds/fruit and yield/
plant indicating the presence of non-allelic interaction,
whereas rest of characters showed non-significant 2
value hence the additive dominance model. For the
rest of the traits where 2 values were significant data
further analyzed for 5 parameter model. All the 5
components were found significant hence this model
was adequate for fruit diameter. For the fruit diameter,
dominance and dominance dominance component
were important whereas, rest of the character were not
significant. Dominance Dominance component was
in general, greater in magnitude than dominance
component for all the crosses. Duplicate type of epistasis
were observed for days to first picking, plant height,
fruit length, fruit diameter and fruit weight in this situation
reciprocal recurrent selection is likely to be useful for
the effective utilization of both type additive and non
additive generation simultaneously. The simple additive-
dominance model revealed the goodness of fit in only 6
characters. The findings were in agreement with
Hayman and Mather (1955) who also reported the
significant deviation from zero indicates the inadequacy
 Vegetable Science, Vol. 41, January - June 2014 39

Table 1: Best fit, estimate of gene effects from analysis of generation means for eleven traits in bitter gourd cross Gy333
DRAR-1
Type of
Traits M D H I l X2 Epistasis
Days to First
3Parameter 41.83** 0.25 -3.81** 0.54 -1.20** 0.41 5.61 C
Flower Anthesis
3 Parameter 60.04** 0.26 -1.25** 0.37 0.17 0.47 8.50*
Days to First
5 Parameter 61.19** 0.66 -1.30 ** 0.37 -3.06 2.94 -1.89 ** 0.58 1.88 2.43
Picking
Best fit 60.31** 0.18 -1.30** 0.37 -1.01 0.42 2.60NS D
3 Parameter 2.31** 0.4 -0.25** 0.04 -0.003 0.06 20.93**
Plant Height 5 Parameter 3.50** 1.21 -0.12**.05 -3.33 3.63 -1.36 1.21 2.07 2.42
Best fit 2.46** 0.48 -0.12 ** 0.05 -0.22** 0.08 -0.32 ** 0.07 0.73NS C
Internodal
3 Parameter 6.97**0.05 -0.61**0.10 0.57**0.06 4.00NS D
Length
Branches/Plant 3 Parameter 11.61**0.17 0.630.39 0.430.34 7.59* C
Fruits/Plant 3 Parameter 30.16**0.72 11.91**0.80 -13.19**0.84 4.16NS D
3Parameter 17.37 ** 0.24 -4.22** 0.28 2.45** 0.42 27.61**
Fruit Length 5Parameter 15.55 ** 0.66 -3.79 ** 0.32 11.94 **2.40 1.22 0.74 -8.43** 1.87
Best fit 16.53**0.29 -3.61**0.30 8.60**1.30 -6.07** 1.22 2.75NS D
3 Parameter 5.25** 0.08 -0.82** 0.14 0.52** 0.15 25.58*
Fruit Diameter 5 Parameter 5.84** 0.17 -0.87 ** 0.14 -2.11** 0.88 -1.10** 0.22 1.99 ** 0.75
Best fit 5.54** 0.05 -0.86 ** 0.13 -0.79 ** 0.14 8.09* C
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3 Parameter 52.43** 0.61 -9.89 ** 0.70 7.29 ** 1.04 7.39*

Fruit Weight 5 Parameter 58.26** 2.35 -9.75** 0.70 -15.38 10.97 -6.53** 2.45 16.70 8.98
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C
Best fit 55.24 ** 0.95 -9.75** 0.70 -3.51** 1.18 4.26**3.19 1.96NS
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Seeds/Fruit Best fit 19.32** 0.30 1.89 ** 0.49 -1.13** 0.54 3.60NS C
Yield/Plant Best fit 1823.73**56.87 689.80**65.32 -815.90**62.8 3.10NS D
C= Complementary, D= Duplicate, NS, *, ** non-significant, significant at p<0.01, <0.05, respectively

of additive-dominance model a presence of epistasis and fruit diameter, fruit weight and yield/ plant in cross
could be tested out. Gy333 DRAR-1, whereas days to first flower anthesis,
plant height, fruit/plant and days to first picking showed
Cross Gy323 DRAR-1 showed significant epistatic
duplicate type epistasis for cross Gy323 DRAR-1.
gene effects. 2 value was significant for all traits except
Duplicate type of epistasis was observed for majority
yield/plant (Table 2). Both additive and dominance
of the traits indicating that heterosis based on dominance
components were s ignific ant for most of tra its.
and dominance dominance interaction gene effect
However, the magnitude of dominance was higher than
cannot be exploited. In such a situation, intermating or
additive indic ating importance of domina nc e
biparental mating between selected plants from early
components . The additive a dditive
segregating generations could help in improving traits
epistaticcomponents were significant for seeds/fruit and
fruit weight, being negative significant for remain all (Shahiet al., 2005).
traits. Therefore this trait was controlled by both additive Most of the traits examine exhibited the combined
and dominant gene actions. influence of substantial dominance and epistatic effects
except for primary branch number and fruit number/
The 3 parameter were found significant along with non-
plant, which were mainly controlled by additive factors
significant 2 value indicating the adequacy of the model
(two to four). Days to first flower anthesis, branches/
for days to first flower anthesis, internodal length, fruit/
plant and seeds/fruit in crossGy333 DRAR-1, whereas
plant, seeds/fruit and yield/plant and these traits showed
fruit length, fruit diameter and seeds/fruit in cross Gy323
duplicate type of epistasis, so dominant component was
DRAR-1 showed complementary epistasis. It is
important. Additive gene action was involved for days
difficult to exploit them due to duplicate type of
to first flower anthesis in bitter gourd, whereas fruit
epistasis. The development of early flowering genotypes
diameter and fruit length both additive and non-additive
possessing desirable fruit weight characteristics (i.e. high
gene action were predominant. Singh et al. (2000)
fruit weight per plant and average weight per fruit),
reported that components of additive gene effects were
however, will likely be complicated by inherent of
significant for fruit length, days to first harvest, fruit
gynoecious in dominance and epistatic effects.
weight and number of fruit/vine and additive and non-
additive gene action was involved in the character In the present study 3 generation (P1, P2 & F1) and 5
expression. generation (P1, P2, F1, F2 & F3) of 2 cross combinations
have been analyzed for 11 characters and the gene
Duplicate type of epistasis was found for fruit length,
effects we re e stimated through pa rtitioning the
 40 Shukla et al. : Genetic analysis of yield and yield attributing traits in bitter gourd

Table 2: Best fit, estimate of gene effects from analysis of generation means for eleven traits in bitter gourd cross Gy323
DRAR-1
Type of
Traits M d h i l X2
Epistasis
3Parameter 39.45 0.30 -1.28 0.42 0.19 0.66 8.8*
Days to First
5Parameter 41.05 **0.63 -1.51** 0.50 -8.35** 3.00 -1.94**0.81 7.39 ** 2.61
Flower Anthesis
Best fit 39.39** 0.24 -1.31** 0.40 0.51 0.63 8.8*
3Parameter 57.99** 0.38 -1.24** 0.55 0.69 0.79 11.34**
Days to First
5 Parameter 59.34** 0.73 -2.70** 0.73 -3.99 3.31 -3.24** 1.03 3.08 2.87
Picking
Best fit 58.47**0.22 -2.70**0.72 -2.37**0.75 1.55NS C
3Parameter 2.18** 0.01 -0.20 ** 0.04 0.002 0.02 46.22**
Plant Height 5Parameter 2.25** 0.02 -0.14** 0.04 -0.29** 0.13 -0.31 ** 0.05 0.22 ** 0.11
Best fit 2.19** 0.01 -0.14** 0.04 -0.25 ** 0.04 6.34* C
Internodal Length Best fit 6.01** 0.06 -0.62 ** 0.08 -0.76** 0.21 2.91NS D
3Parameter 9.49** 0.35 2.13 ** 0.48 -3.22** 0.54 10.74**
Branches/Plant 5Parameter 11.31** 0.86 2.40** 0.53 -13.61** 3.66 -1.31 1.01 8.91 ** 2.95
Best fit 10.36** 0.45 2.59** 0.51 -9.91** 2.29 6.17 ** 2.05 1.68NS D
3Parameter 31.08** 0.65 11.98 **0.70 -17.39 **0.85 7.04*
Fruits/Plant
5Parameter 39.23 ** 3.2 12.20 **0.71 -47.44**13.57 -8.63** 3.28 21.88 ** 10.34
Best fit 31.08 ** 0.65 11.98** 0.70 -17.39 **0.84 7.03* D
3Parameter 18.76** 0.26 -3.95** 0.31 2.96** 0.58 38.75**
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Fruit Length 5Parameter 20.65 ** 0.63 -3.14 ** 0.34 -0.16 2.66 -3.23** 0.72 -0.23 2.30
Best fit 20.52** 0.19 -3.14** 0.34 -3.10 ** 0.39 0.27NS C
3Parameter 5.34** 0.82 -0.61** 0.13 -0.05 0.17 10.47**
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Fruit Diameter 5Parameter 5.56 ** 0.17 -0.58** 0.13 -0.44 0.69 -0.53 ** 0.21 -0.01 0.60
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Best fit 5.57 ** 0.11 -0.58** 0.13 -0.45** 0.21 -0.54** 0.17 0 C
3Parameter 53.04** 0.50 -6.70 ** 0.57 -7.51** 0.79 11.38**
Fruit Weight 5Parameter 49.07** 1.66 -7.35 ** 0.61 3.58 6.97 5.06 ** 1.76 -6.79 5.57
Best fit 49.85 ** 0.66 -7.35 ** 0.61 4.28** 0.90 -3.97** 0.94 0.26NS C
3Parameter 13.51** 0.27 1.60** 0.67 2.90** 0.65 68.49**
Seeds/Fruit 5Parameter 12.74** 0.49 0.50 0.69 3.51 2.58 5.96 ** 0.84 1.27 2.33
Best fit 12.53 ** 0.30 4.87 ** 0.70 6.28 ** 0.73 0.83NS C
Yield/Plant Best fit 1639.37** 41.14 489.90**47.11 -1011.65**48.45 3.44 C
C= Complementary, D= Duplicate, NS, *, ** non-significant, significant at p<0.01, <0.05, respectively

generation means into different components. In the event
of inadequacy of the model the data was refitted for
both parameter models. For the characters where all
the 3 and 5 components were not significant in the both
paramete r model the elimination of the gene tic
component was also done to increase the precision of
the results.
Positive additive additive is indicating the possibility
of obta ining transgre ss ive se gregants in la te r
generations, whereas dominance dominance gene
effects indicated a duplicate epistasis which will be
undesirable for selection and genetic improvement.
Similarly, dominance in the direction of higher yield,
a ss oc ia te d with a negative domina nc e
domina nceinte raction will be undesirable when
improvement in yield is the gynoecious. Gene action
and empirical estimates of genetic parameters governing
trait expression have been useful in developing breeding
strategies for incorporating gynoecious for high yielding
in cucumber (Fazio et al.,1994 and Ram et al., 1997).
The additive dominance and dominance dominance
interaction effects were noted in most of the crosses
for all the traits. The complementary type of epistasis
was noted in all the traits except internodal length,
branches/plant and fruit/plant for Gy323 DRAR-1 and
days to first picking, internodal length, fruit/plant, fruit
length and yield/plant for Gy333 DRAR-1. These traits
showed duplicate type of epistasis for the above crosses.
Results suggest that introgression of yield-related from
gynoecious into monoecious may aid in the development
of high-yielding cultivars with gynoecious. Strategies
for the development of gynoecious can be determined.
These strategies will likely incorporate the use of both
molecular and conventional breeding (reciprocal
recurrent selection) for designing an optimal approach
for increasing yield in bitter melon.
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