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Breeding Okra for Higher Productivity and Yellow Vein Mosaic

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Breeding Okra for Higher Productivity

and Yellow Vein Mosaic Tolerance
a a b
Sibsankar Das , Arup Chattopadhyay , Subrata Dutta ,
a a
Sankhendu Bikash Chattopadhyay & Pranab Hazra
a
Department of Vegetable Crops, Faculty of Horticulture, Bidhan
Chandra Krishi Viswavidyalaya, Mohanpur, Nadia, West Bengal, India
b
Department of Plant Pathology, Faculty of Agriculture, Bidhan
Chandra Krishi Viswavidyalaya, Mohanpur, Nadia, West Bengal, India
Version of record first published: 10 Jan 2013.

To cite this article: Sibsankar Das , Arup Chattopadhyay , Subrata Dutta , Sankhendu Bikash
Chattopadhyay & Pranab Hazra (2013): Breeding Okra for Higher Productivity and Yellow Vein Mosaic
Tolerance, International Journal of Vegetable Science, 19:1, 58-77

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 International Journal of Vegetable Science, 19:58–77, 2013
Copyright © Taylor & Francis Group, LLC
ISSN: 1931-5260 print / 1931-5279 online
DOI: 10.1080/19315260.2012.675024

Breeding Okra for Higher

Productivity and Yellow Vein
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Mosaic Tolerance
Sibsankar Das,1 Arup Chattopadhyay,1 Subrata Dutta,2
Sankhendu Bikash Chattopadhyay,1 and Pranab Hazra1
1
Department of Vegetable Crops, Faculty of Horticulture, Bidhan Chandra Krishi
Viswavidyalaya, Mohanpur, Nadia, West Bengal, India
2
Department of Plant Pathology, Faculty of Agriculture, Bidhan Chandra Krishi
Viswavidyalaya, Mohanpur, Nadia, West Bengal, India
Improving yield and ensuring its sustainability under adverse conditions through
resistant hybrids is the major objective of heterosis breeding in okra (Abelmoschus escu-
lentus L. Moench). A study was undertaken to determine gene action, heterobeltiosis,
and combining ability for 15 characters through a Line × Tester mating design in okra.
Nonadditive gene action controlled all characters studied. Crosses showing high spe-
cific combining ability (sca) and yield involved parents showing high general combining
ability for plant height, number of nodes on the main stem, number of leaves per plant,
fruit diameter, fruit weight, and number of fruits per plant. Parental lines and hybrids
were categorized according to disease reaction. The hybrids ‘VNR Green’ × ‘Shagun’ and
‘Barsha Laxmi’ × ‘Parbhani Kranti’ were selected on the basis of their mean (per se)
performance, heterosis manifested in them, and sca effects. These hybrids could be used
commercially due to high yield and low percentage disease index values for yellow vein
mosaic virus (YVMV) disease.

Keywords Abelmoschus esculentus, Combining ability, Gene action, Heterosis, India.

Okra (Abelmoschus esculentus L. Moench) is primarily a warm season annual

vegetable crop grown in tropical and subtropical parts of the world. The crop
can express variability in plant and fruit type and landraces, and primitive
types can be found with poor yields and varying degrees of tolerance to stresses
(B. S. Dhankar and Mishra, 2005).

The authors gratefully acknowledge Dr. Bijendra Singh, Project Coordinator, AICRP on
Vegetable Crops, ICAR, New Delhi, India, for providing financial assistance to carry out
this experiment.
Address correspondence to Arup Chattopadhyay, Department of Vegetable Crops,
Faculty of Horticulture, Bidhan Chandra Krishi Viswavidyalaya, Mohanpur 741252,
Nadia, West Bengal, India. E-mail: chattopadhyay.arup@gmail.com
 Breeding Okra 59

Despite its high nutritive value and acceptability among growers and con-
sumers, average productivity of okra in India is not as good as that of other
okra-growing countries. One of the major constraints in okra production is
yellow vein mosaic virus (YVMV) disease, which can occur at all crop growth
stages (Capoor and Verma, 1950; Kulkarni, 1924; Uppal et al. 1942; Verma,
1952). This disease is caused by a complex consisting of the monopartite bego-
movirus and a small satellite DNA β component (Jose and Usha, 2003). The
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virus is neither sap nor seed transmissible. In nature, transmission occurs

through the whitefly (Bemisia tabaci Genn.) insect vector. Infection of 100%
of plants in a field is usual and yield losses range from 50% to 94% depending
on stage of crop growth at which infection occurs (Sastry and Singh, 1974).
Only a few attempts have been made to study genetics of okra
resistance/susceptibility to YVMV. There is no definite agreement for how to
achieve inheritance of resistance to YVMV. H. B. Singh et al. (1962) found that
two recessive alleles at two loci conferred resistance in intervarietal crosses of
okra. Thakur (1976) implicated two complementary genes that govern resis-
tance to YVMV under natural epiphytotic conditions. Dhillon (1978) agreed
with these results and stated that additive component of variance was predom-
inant compared to dominant gene effects. They also concluded that the genes
governing YVMV resistance were influenced by environmental conditions and
were temperature sensitive. There might be a possibility of polygenic control
of YVMV in okra. Arumugam and Muthukrishnan (1980) and Jambhale and
Nerkar (1981) revealed that resistance to YVMV was controlled by a single
dominant gene. Sharma and Dhillon (1983) suggested that there are two com-
plementary genes governing resistance to YVMV. Ali et al. (2000) hypothesized
that tolerance to YVMV is quantitative, with possibly two major factors, and
dependent on gene dosage with incompletely dominant gene action. Vashisht
et al. (2001) indicated that there is complex genetic control of resistance to
YVMV and reported that inheritance of YVMV was governed by epistatic gene
action. S. K. Dhankar et al. (2005) confused the issue by reporting that inher-
itance of resistance to YVMV is under control of two complementary genes
following Mendelian segregation.
Okra can be cross-pollinated and can be exploited for production of F1
hybrids. Heterosis leads to increase in yield, reproductive ability, adaptability,
disease and insect resistance, general vigor, and quality. Yield improvement
in okra of 49% to 65% occurs through heterosis breeding (Joshi et al., 1958;
Kumar and Sreeparvathy, 2010; Partap et al., 1981). In okra, fruit number is
the most effective factor contributing to yield, followed by plant height, fruit
weight and size, and earliness (B. S. Dhankar and Mishra, 2005). The floral
biology of okra enables easy emasculation and pollination and the plant pro-
duces a large number of seeds inform a single pollination, which makes the
commercial exploitation of heterosis possible in okra even with hand emascu-
lation and pollination. Exploitation of hybrid vigor for an economic character
 60 S. Das et al.

in a crop is successfully utilized from a specific cross-combination of parents

determined from the study of combining ability analysis. Information on the
extent and nature of gene action is essential for framing a sound strategy for
combination breeding to improve yield and other characters. Several reports on
the extent of heterosis and combining ability in okra are available, but ample
utilization of this information for commercial production of okra hybrids is
lacking compared to other common vegetables in India. This project was under-
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taken to study combining ability through Line × Tester analysis to determine

the nature and magnitude of heterosis for yield component characters, quality
attributes, and YVMV tolerance; identify good general combiners; determine
the breeding strategies for genetic improvement; and assess disease tolerance
of parental lines and hybrids against YVMV disease.

MATERIALS AND METHODS

The investigation was carried out during spring–summer season beginning in
Feb. 2009 and continued up to Sept. 2010. The location of the experiment was
the Research Farm of Bidhan Chandra Krishi Viswavidyalaya, Kalyani, Nadia,
West Bengal, India, situated at 23.5◦ N latitude and 89◦ E longitude with a
mean sea level of 9.75 m.

Development of F1 Hybrids and Field Trials

Seed of the YVMV-tolerant lines ‘Barsha Laxmi’, ‘Deb-401’, and ‘VNR
Green’ and the testers ‘Parbhani Kranti’, ‘Bengal IMP’, and ‘Shagun’, which
have high yield and good quality attributes, were used. Fertilizer at the rate
of 60N–60P–60K kg·ha−1 was applied preplant to the soil. Nitrogen was from
urea, P was from single superphosphate, and K was from muriate of potash.
Additional nitrogen at 60 kg·ha−1 was applied in two equal splits at 30 and
55 days after planting. Seed of parental lines, after treatment with Thiram
(Devidayal Agrochemicals Limited, Mumbai, India) (3 g·kg−1 of seed), were
sown during the first week of Feb. 2009 in 2.5-cm-deep furrows with 45-cm
spacing between rows and 30 cm between plants in plots that were 2.25 ×
2.4 m. Management was as per Chattopadhyay et al. (2007).
Crossing was carried out during full bloom. Flower buds measuring 3–5 cm
long and 1.5–2 cm in diameter, which would likely to open the next morn-
ing, were chosen for emasculation in the afternoon hours between 4 and 6 PM.
A circular cut was made around the cone, the fused calyx including the epica-
lyx, and the closely packed overlapping petals with a sharp blade about 1 cm
above the base. Care was taken so that the staminal column and gynoecium
were not injured. After the incision was made, the cone-shaped epicalyx was
pulled off along with the petals, exposing the staminal tube and stigma. The
anthers were removed from the staminal column with forceps. After emascu-
lation, instead of bagging, the cap-like epicalyx along with petals was fixed
 Breeding Okra 61

over the stigma to prevent unwanted pollination. Forceps were sterilized in

absolute alcohol before and after each operation to prevent pollen contamina-
tion. The next morning, the cap of epicalyx and petals was removed from the
emasculated flower bud and it was pollinated between 7 and 8 AM with pollen
from the intended male parent and again covered with the same cap to prevent
outcrossing. The cap was removed 2–3 days after pollination. Each flower bud
after emasculation and pollination was tagged and labeled indicating parents
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involved in cross and the date of crossing. Selfing of each parental line was
also done to maintain genetic purity. Matured seed of 35-day-old pods from
the parents and hybrids were collected, sun-dried, and stored separately in
desiccators for sowing the next season.
In 2010, seeds of six parental lines and nine hybrids were sown during
the first week of June as described previously in a randomized complete block
design with three replications. Plants were spaced at 60 × 30 cm with 60 plants
in each replication in plots 3.6 × 3.0 m. No insecticides were used in order to
build up a reasonable white fly (Bemicia tabaci Gen.) population, the vector
of YVMV. The susceptible variety ‘Pusa Sawani’ was planted and maintained
around the field to ensure sufficient virus inocula. Observation on plant height
at flowering, number of lateral branches per plant, number of nodes on the
main stem, number of leaves per plant, days to first flowering, nodes at first
flowering, fruit length, fruit diameter, fruit weight, number of fruit per plant,
number of seed per fruit, and fruit yield per plant were recorded from 15 ran-
domly selected plants from each entry. Ascorbic acid (Ranganna, 1986) and
crude protein contents (nitrogen content by micro-Kjeldahl method × 6.25) of
young fruit were also estimated.

Observations on Disease Parameters

Percentage of disease index (PDI) on lines, testers, and F1 s was recorded
at four stages at an interval of 15 days starting from 30 days after sowing
(DAS) to 75 DAS. Vein clearing of any form in the plant was treated as dis-
ease incidence. The PDI was expressed as a percentage from all 60 plants in a
replication using a disease severity scale (0–9) for plants through visual eval-
uation. The disease severity scale for tomato leaf curl virus disease developed
by Banerjee and Kalloo (1987) was adopted as a reference for developing this
scale. In the initial evaluation, no genotype was disease free and on an average
at least 10%–15% of leaf area of the genotypes showed disease symptoms. The
disease severity rating was assessed as follows: 0 = no disease, 1 = up to 15%
of leaf area affected, 3 = 30%–45% of leaf area affected, 5 = 45%–60% of leaf
area affected, and 7 = greater than 60% of leaf area affected.
YVMV disease was rated according to the scale of PDI values, where resis-
tant (R) had a PDI ≤ 10%, moderately resistant (MR) 11 < PDI ≤ 15%,
moderately susceptible (MS) 16 < PDI ≤ 45%, and highly susceptible (HS)
 62 S. Das et al.

had a PDI > 45%. The number of plants infected in each entry was recorded
and the PDI calculated with the following:

Numerical ratings
PDI = × 100
Highest grade of rating × Total number of plants of the entry examined

Statistical Analyses
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Data were analyzed with the Line × Tester model of genetic anal-
ysis (Kempthorne, 1957) using the software SPAR I (Indian Agricultural
Statistical Research Institute, New Delhi, India). Heterosis over better par-
ent (heterobeltiosis) was estimated as a percentage increase or decrease of the
F1 hybrid over its better parent (Hayes et al., 1965) using the formula:

Heterobeltiosis (%) = [F1 − BP/BP] × 100

Significance of better parent heterosis was determined with the t-test (Wynne
et al., 1970).

√
SE (BP) = (2 MSE/r); t-value = Heterobeltiosis/SE (BP)

where BP is the mean of better parent in the cross; MSE is the mean square
error; and r is the number of replications.
Combining ability was analyzed according to Nadarajan and Gunasekaran
(2008) based on Griffing’s (1956) fixed effect model using the formula:

Yij = m + gi + gj + sij + rij + 1/bc ijkl

where i, j = 1, 2, . . . , n; k = 1, 2, . . . , b. l = 1, 2, . . . , c; Yij is the mean of i × j

genotype over k and l; m is the population mean; gi is the general combining
ability (gca) effect of the ith parent; gj is the gca effect of the jth parent; sij is the
specific combining ability (sca) effect; rij is the reciprocal effect; and 1/bc ijkl
is the mean error effect. Data were subjected to analysis of variance (Panse and
Sukhatme, 1984). Heterobeltiosis was computed using the software program
Windowstat 8.0 (Indostat Services 18, Ameerpet, Hyderabad, India).

RESULTS AND DISCUSSION

Nature of Gene Action

The general and specific combining ability are the main criteria of rapid
genetic assay of the test genotypes under Line × Tester analysis. Combining
ability variances were not significant for lines in all of the characters. However,
the variances were significant for testers in days to first flowering and fruit
 Breeding Okra 63

length and for Line × Tester in all characters (Table 1). The gca and sca showed
a wide range of variation for all of the characters studied. Additive variance
(α2 A) and dominance variance (a2 D) can be calculated at F = l (okra is often
a cross-pollinated crop) from gca and sca variances. In this prediction, α2 D =
2α2 gca and α2 H = α2 sca. A general trend of the genetic control of the char-
acters can be ascertained from these estimates of additive and nonadditive
variance components. The relative importance of the additive and nonadditive
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genetic effects for these characters was reflected by the predictability ratio
α2 D/(α2 H + α2 D) as per Baker (1978). The closer this ratio is to unity, the
greater the predictability based on gca alone.
There was a preponderance of nonadditive gene action in the control for
all characters studied; selection will result in no or slow genetic improve-
ment (Table 2). The overwhelming response of nonadditive gene action for
conditioning of plant height (Ezhilarasi and Thangavel, 2011; Ramya et al.,
2010), number of lateral branches per plant (B. S. Dhankar and Dhankar,
2001), number of nodes per plant (Dabhi et al., 2010; Ramya et al., 2010),
days to first flowering (Dabhi et al., 2010; Das et al., 1996), number of nodes
at first flowering (Dabhi et al., 2010), fruit length (Dabhi et al., 2010; Ramya
et al., 2010), fruit diameter (Das et al., 1996; Ramya et al., 2010), fruit weight
(Ezhilarasi and Thangavel, 2011; More and Patil, 1997), number of fruit per
plant (Ezhilarasi and Thangavel, 2011; Ramya et al., 2010), and fruit yield per
plant (More and Patil, 1997; Ramya et al., 2010; B. Singh and Sanwal, 2010;
Solankey and Singh, 2010) have been reported irrespective of the parental
materials used, methods followed, and environments in which experiments
were conducted. Reports by Reddy et al. (1985), Wankhade et al. (1995), Das
et al. (1996), Mandal and Das (1992), Sood and Sharma (2001), and Arora et al.
(2008) emphasized the importance of additive genetic effects for plant height,
number of lateral branches per plant, number of nodes on the main stem, days
to first flowering, nodes at first flowering, number of fruits per plant, fruit
diameter, fruit length, fruit weight, fruit yield per plant, and YVMV incidence.
However, Chandra et al. (1997), while examining the genetic control of num-
ber of seeds per fruit, found that both additive and nonadditive gene effects
were involved. No information is available on the nature of gene action govern-
ing number of leaves per plant, PDI of YVMV disease, and ascorbic acid and
crude protein contents of young fruit in okra. The presence of additive gene
effects for days and nodes to YVMV appearance has been reported (Sharma
and Dhillon, 1983). Successful breeding methods are those that accumulate the
genes to form superior gene constellations interacting in a favorable manner.
The importance of nonadditive gene action for all yield components, qual-
ity traits, and PDI of YVMV in the present study indicates that heterosis
breeding is the best possible option for improving these traits in okra. This is
the common breeding strategy followed for the characters where nonadditive
genetic effects had a great impact on their genetic variation. The disparities
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64
Table 1: Analysis of variance for combining ability.

Mean sum of squares for parents and hybrids for the character
Source of
variation (df) PHa NPBPP NNMS NLPP DFF NFF FL FD FW NFPP NSPF AAC CP FYPP PDI

Replication (2) 3.39 0.00 0.09 0.47 0.10 0.05 0.36 0.01 0.30 1.07 1.58 0.18 0.00 165.70 0.49
Lines (2) 2.88 0.55 0.28 0.23 2.28 0.14 1.97 0.01 0.11 12.23 154.73 7.38 0.01 4,597.65 40.145
Testers (2) 177.29 1.63 6.54 8.38 104.45∗ 2.36 11.00∗∗ 0.07 6.79 19.68 0.79 0.72 0.01 21,490.60 124.83
Line vs. Tester (4) 72.21∗∗ 1.19∗∗ 7.76∗∗ 6.95∗∗ 22.41∗∗ 1.06∗∗ 1.83∗∗ 0.07∗∗ 5.85∗∗ 38.22∗∗ 179.70∗∗ 2.36∗ 0.11∗∗ 29,180.17∗∗ 37.61∗∗
Error (16) 1.15 0.01 0.13 0.45 0.16 0.02 0.21 0.00 0.15 0.34 8.34 0.65 0.004 177.22 1.75

a PH = plant height, NPBPP = number of lateral branches per plant, NNMS = number of nodes on the main stem, NLPP = number of leaves per plant,

DFF = days to first flowering, NFF = nodes at first flowering, FL = fruit length, FD = fruit diameter, FW = fruit weight, NFPP = number of fruits per plant,
NSPF = number of seeds per fruit, AAC = ascorbic acid content, CP = crude protein, FYPP = fruit yield per plant, PDI = percentage disease index at
75 DAS.
∗, ∗∗ Significant at 5% or 1% levels, two-tailed t-test.
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Table 2: Estimates of component of variancea .

Component of
genetic variance PH NPBPP NNMS NLPP DFF NFF FL FD FW NFPP NSPF AAC CP FYPP PDI

α2 GCA 0.496 0.002 0.120 0.073 0.859 0.005 0.129 0.0008 0.066 0.618 2.831 0.047 0.003 448.223 1.246
α2 D (2 α2 GCA) 0.992 0.004 0.240 0.146 1.718 0.010 0.258 0.0016 0.132 1.236 5.662 0.094 0.006 896.446 2.492
α2 H (α2 SCA) 26.66 0.377 1.81 1.727 12.575 0.379 1.316 0.0176 1.500 8.915 40.127 0.850 0.020 6978.309 19.434
σ 2 H/σ 2 D 26.88 94.25 7.54 11.83 7.32 37.90 5.10 11.00 11.36 7.21 7.09 9.04 3.33 7.78 7.80
Predictability Ratio 0.035 0.010 0.117 0.078 0.120 0.025 0.164 0.083 0.080 0.121 0.123 0.010 0.230 0.114 0.113
α2 D/(α2 H + α2 D)

a PH = plant height, NPBPP = number of lateral branches per plant, NNMS = number of nodes on the main stem, NLPP = number of leaves per plant,

DFF = days to first flowering, NFF = nodes at first flowering, FL = fruit length, FD = fruit diameter, FW = fruit weight, NFPP = number of fruits per plant,
NSPF = number of seeds per fruit, AAC = ascorbic acid content, CP = crude protein, FYPP = fruit yield per plant, PDI = percentage disease index at
75 DAS.

65
 66 S. Das et al.

in interpretation between the present study and previous reports might be

due to differences in parental materials used and the environment under
which the trial was conducted because estimates of gene effects change with
environments and genotypes.

gca and Mean Performance

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After the overall picture of gca effects was assessed, it appeared that the
parents differed in their gca (Table 3). Among lines, the highest significant
and positive gca effects were for ‘Barsha Laxmi’ for number of lateral branches
per plant, days to first flowering, nodes at first flowering, number of fruits
per plant, number of seeds per fruit, ascorbic acid content of fruit, crude pro-
tein content of fruit, and fruit yield per plant. Significant gca effects were
also recorded by ‘VNR Green’ for fruit yield per plant, number of fruit per
plant, and fruit length. Among testers, ‘Parbhani Kranti’ exhibited positive
gca effects for plant height, fruit length, fruit diameter, fruit weight, number
of fruits per plant, and fruit yield per plant. Negative significant gca effects for
days to first flowering and nodes at first flowering were for ‘Parbhani Kranti’
and ‘Shagun’. The highest mean performance for fruit yield per plant was for
‘Deb-401’, and the lowest mean for days to 50% flowering was for ‘Parbhani
Kranti’.
When assessed for general combining ability, the parents ‘Barsha Laxmi’
and ‘Parbhani Kranti’ were good combiners for fruit yield and quality-related
traits. Because a high gca effect is related to additive and Additive × Additive
interaction and represents fixable components of genetic variance, ‘Barsha
Laxmi’ and ‘Parbhani Kranti’ could be used in breeding for high yield and
better fruit quality. This result supports observations of Pathak et al. (1998),
Shukla et al. (1989), and Sood and Kalia (2001), who found that ‘Parbhani
Kranti’ was a good general combiner when tested in other environments.
Significant negative gca effects of days to 50% flowering and nodes at first
flowering for testers ‘Parbhani Kranti’ and ‘Shagun’ indicate that these have
additive genetic effects for decreasing these traits.

Heterobeltiosis and sca

For plant height, the maximum significantly positive heterobeltiosis
occurred in the ‘VNR Green’ × ‘Shagun’ cross followed by ‘Barsha Laxmi’ ×
‘Parbhani Kranti’ (Table 4). Heterobeltiosis for this character ranged from
−2.97% to 15.53%. The best hybrid based on mean performance was ‘Barsha
Laxmi’ × ‘Parbhani Kranti’, displaying heterosis of 14.69% over the better
parent. Heterosis for plant height within this magnitude supports the find-
ings of Jaiprakashnarayan et al. (2008), Khanpara et al. (2009), B. Singh
and Sanwal (2010), and Akhtar et al. (2010) for different genotypes and
environments.
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Table 3: Estimates of general combining ability effects in six parents over nine F1 sa .

Parents PH NPBPP NNMS NLPP DFF NFF FL FD FW NFPP NSPF AAC CP FYPP PDI

Barsha −0.64 0.18∗∗ −0.05 −0.14 0.58∗∗ 0.15∗∗ −0.38∗ −0.04 −0.10 0.81∗∗ 2.54∗ 0.78∗∗ 0.05∗ 14.53∗∗ −2.18∗∗
Laxmi (56.78)b (3.79) (24.55) (23.76) (44.03) (5.27) (12.30) (1.80) (18.65) (22.88) (57.13) (15.06) (2.04) (412.62) (8.72)
Deb-401 0.43 −0.28∗∗ 0.20 −0.04 −0.28∗ −0.07 −0.14 −0.004 −0.02 −1.34∗∗ 2.25∗ −1.00∗∗ −0.03 −26.04∗∗ 2.04∗∗
(45.73) (4.17) (23.37) (22.88) (44.38) (5.18) (12.28) (1.67) (20.25) (24.76) (57.45) (13.88) (1.90) (487.93) (7.74)
VNR 0.21 0.10∗ −0.15 0.18 −0.30∗ 0.07 0.52∗∗ 0.04 0.12 0.53∗ −4.78∗∗ 0.22 −0.02 11.51∗ 0.14
Green (74.07) (3.28) (26.13) (25.98) (47.02) (5.42) (15.96) (1.84) (17.37) (19.30) (56.96) (14.27) (1.84) (324.16) (10.29)
Parbhani 3.37∗∗ −0.27∗∗ 0.09 0.40 −2.12∗∗ −0.25∗∗ 1.13∗∗ 0.10∗∗ 0.99∗∗ 1.64∗∗ −0.23 0.33 −0.04 54.49∗∗ −3.81∗∗
Kranti (75.69) (1.86) (26.14) (28.42) (43.13) (5.22) (14.83) (1.84) (18.92) (20.96) (69.00) (13.93) (1.98) (383.29) (16.68)
Bengal −5.03∗∗ 0.49∗∗ −0.90∗∗ −1.10∗∗ 3.93∗∗ 0.59∗∗ −1.08∗∗ −0.05∗ −6.65∗∗ −1.23∗∗ −0.10 −0.15 0.03 −39.93∗∗ 3.63∗∗
IMP (75.67) (3.33) (26.03) (27.53) (45.20) (5.45) (13.16) (1.90) (18.14) (17.01) (62.82) (13.04) (1.92) (301.39) (23.36)
Shagun 1.66∗∗ −0.23∗∗ 0.80∗∗ 0.70∗∗ −1.81∗∗ −0.34∗∗ −0.05 −0.04 −0.34∗ −0.40 0.34 −0.18 0.004 −14.56∗∗ 0.18
(72.89) (2.22) (26.40) (28.97) (47.17) (5.31) (12.55) (1.84) (20.41) (19.93) (70.67) (14.04) (2.07) (397.06) (19.37)
SE (gi ) 0.36 0.04 0.12 0.22 0.13 0.05 0.15 0.02 0.13 0.20 0.96 0.27 0.02 4.44 0.44

a PH = plant height, NPBPP = number of lateral branches per plant, NNMS = number of node on the main stem, NLPP = number of leaves per plant,
DFF = days to first flowering, NFF = nodes at first flowering, FL = fruit length, FD = fruit diameter, FW = fruit weight, NFPP = number of fruits per plant,
NSPF = number of seeds per fruit, AAC = ascorbic acid content, CP = crude protein, FYPP = fruit yield per plant, PDI = percentage disease index at
75 DAS.
b Values in parentheses indicate mean performance.
∗, ∗∗ Significant at 5% and 1% levels, two-tailed t-test.

67
 68 S. Das et al.

Table 4: Heterosis over better parent and their corresponding specific combining
ability.

Heterobeltiosis Specific
Characters Better crosses (%) combining effects

Plant height VNR Green × Shagun 15.53∗∗ 4.71∗∗ (85.58)a

Barsha Laxmi × Parbhani Kranti 14.69∗∗ 5.08∗∗ (86.81)
Deb-401 × Shagun 11.89∗∗ −0.15 (81.56)
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Numbers of VNR Green × Shagun 10.35∗∗ 1.83∗∗ (29.14)

nodes on the Deb-401 × Parbhani Kranti 7.36∗∗ 0.35 (28.06)
main stem Barsha Laxmi × Parbhani Kranti 4.78∗∗ 0.69∗∗ (27.39)
Number of VNR Green × Shagun 4.51∗∗ 1.26∗∗ (30.27)
leaves per Barsha Laxmi × Parbhani Kranti 3.78∗∗ 1.09∗∗ (29.49)
plant Deb-401 × Parbhani Kranti 1.43∗ 0.65 (28.83)
Days to first Barsha Laxmi × Parbhani Kranti −10.75∗∗ −2.14∗∗ (38.49)
flowering Deb-401 × Parbhani Kranti −10.87∗∗ 0.55∗ (38.44)
VNR Green × Shagun −17.72∗∗ −1.37∗∗ (38.69)
Nodes at first Deb-401 × Shagun −21.43∗∗ 0.45∗∗ (4.07)
flowering Barsha Laxmi × Parbhani Kranti −22.60∗∗ −0.53∗∗ (4.04)
VNR Green × Shagun −24.48∗∗ −0.25∗∗ (4.01)
Fruit length Deb-401 × Shagun 12.22∗∗ −0.43 (14.08)
Barsha Laxmi × Shagun 10.34∗∗ −0.61∗ (13.84)
Deb-401 × Parbhani Kranti 8.66∗∗ 0.24 (16.11)
Fruit diameter Deb-401 × Parbhani Kranti 9.62∗∗ 0.08 (2.01)
VNR Green × Shagun 6.09∗∗ 0.20∗∗ (2.03)
Barsha Laxmi × Parbhani Kranti 4.54∗∗ 0.01 (1.92)
Fruit weight Barsha Laxmi × Parbhani Kranti 13.20∗∗ 0.60∗ (21.41)
Deb-401 × Parbhani Kranti 6.54∗∗ 0.43 (21.57)
VNR Green × Shagun 5.34∗∗ 1.79∗∗ (21.50)
Number of fruit VNR Green × Shagun 30.57∗∗ 3.65∗∗ (26.03)
per plant VNR Green × Bengal IMP 14.54∗∗ −2.40∗∗ (22.11)
Barsha Laxmi × Parbhani Kranti 12.25∗∗ 0.99∗∗ (25.68)
Number of seed Barsha Laxmi x Bengal IMP 13.30∗∗ 2.71 (71.18)
per fruit Deb-401 × Bengal IMP 9.37∗∗ 1.57 (68.71)
Ascorbic acid Deb-401 × Parbhani Kranti 4.55∗∗ 0.09 (14.57)
content VNR Green × Shagun 2.29∗∗ 0.46 (14.59)
Crude protein Deb-401 × Bengal IMP 3.47∗∗ 0.07 (1.99)
Barsha Laxmi × Parbhani Kranti 1.96∗∗ 0.16∗∗ (2.08)
Fruit yield per VNR Green × Shagun 36.27∗∗ 112.30∗∗ (541.08)
plant Barsha Laxmi × Parbhani Kranti 29.69∗ 34.30∗∗ (535.14)
VNR Green × Bengal IMP 24.12∗ −63.24∗∗ (402.37)
PDI at 75 DAS VNR Green × Shagun −12.95∗∗ −3.44∗∗ (9.11)
Barsha Laxmi × Parbhani Kranti −11.46∗∗ −3.30∗∗ (7.72)
Deb-401 × Parbhani Kranti −6.84∗∗ −3.24∗∗ (7.21)
∗, ∗∗ Significantat P ≤ 0.05 and 0.01 levels, respectively, critical difference test.
a Values in parentheses indicate mean performance.

None of the hybrids exhibited significant positive heterosis for num-

ber of lateral branches per plant over the better parent. This supports the
observations of Kumar and Sreeparvathy (2010) for different genotypes.
Four of nine hybrids exhibited significant positive heterosis over the better
parent for number of nodes on the main stem. The maximum heterobeltiosis
was for ‘VNR Green’ × ‘Shagun’. Heterosis for this character supports the
reports of Jaiprakashnarayan et al. (2008) and Khanpara et al. (2009) for
different genotypes and environments.
 Breeding Okra 69

Heterobeltiosis for number of leaves per plant ranged from −6.96% to

4.51%. The range of heterobeltiosis of the present study was within the range
reported by Jaiprakashnarayan et al. (2008).
For days to first flowering, six of nine crosses exhibited significant neg-
ative heterosis over the better parent, with the maximum occurring in the
‘VNR Green’ × ‘Shagun’ cross followed by the ‘Deb-401’ × ‘Parbhani Kranti’
cross. Heterobeltiosis for this character ranged from −17.72% to 7.26%.
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Selection of hybrids with negative heterosis over their better parents for this
character may be useful for developing early yielding hybrids. The positive
heterosis for this character indicated that there is no possibility for selection
of early yielding plant type. Heterosis for earliness in okra occurred most
often when one of the parents was early. The remaining crosses displayed
delayed maturity. The lateness can be attributed to the strong influence of
male parents that were late. On the basis of mean performance, ‘Parbhani
Kranti’ was one of the earliest parents and it was involved as a parent
in best hybrid, ‘Barsha Laxmi’ × ‘Parbhani Kranti’, took nearly 39 days in
flowering, and exhibited better effects in other promising hybrids. Heterosis
for this character by Line × Tester analysis supports the findings of Rewale
et al. (2003) and Jaiprakashnarayan et al. (2008) for different genotypes and
environments.
Seven of nine hybrids exhibited significant negative heterosis for node
number at first flowering over the better parent. The range of heterobeltiosis
for this character was −24.48% to 6.95% and supports the observations of
Jaiprakashnarayan et al. (2008) for different genotypes.
The maximum positive heterobeltiosis for fruit length was in the ‘Deb-
401’ × ‘Shagun’ cross followed by the ‘Barsha Laxmi’ × ‘Shagun’ cross.
Heterobeltiosis ranged from −13.49% to 12.22%. The best hybrid on the
basis of mean performance was the ‘VNR Green’ × ‘Shagun’ cross with a
heterosis of 2.13% over the better parent. Heterosis for this character by
Line × Tester analysis supports the observations of Pathak et al. (1998) and
Jaiprakashnarayan et al. (2008) for different parents and environments.
Of nine hybrids, four exhibited significant positive heterosis for fruit diam-
eter over the better parent. The range of heterobeltiosis was −13.22% to
9.62%. Heterosis for this character was similar to that reported by Shukla and
Gautam (1990) for different parents.
The maximum positive significant heterobeltiosis for fruit weight occurred
in the ‘Barsha Laxmi’ × ‘Parbhani Kranti’ cross, followed by the ‘Deb-
401’ × ‘Parbhani Kranti’ cross. The range of heterobeltiosis was −9.62% to
13.20%. The best hybrid on the basis of mean performance was the ‘Deb-
401’ × ‘Parbhani Kranti’ cross with a heterosis of 6.54% over the better
parent. Heterosis for this character by Line × Tester analysis supports the
observations of Jaiprakashnarayan et al. (2008) for dissimilar parents and
environments.
 70 S. Das et al.

Four out of nine hybrids exhibited significant positive heterosis for num-
ber of fruits per plant over the better parent. The range of heterobeltiosis
was −23.91% to 30.57%. The maximum positive heterobeltiosis was the ‘VNR
Green’ × ‘Shagun’ cross followed by the ‘VNR Green’ × ‘Bengal IMP’ cross.
The best hybrid on the basis of mean performance was the ‘VNR Green’ ×
‘Shagun’ cross with a heterosis of 30.57% over the better parent. Heterosis for
this character by Line × Tester analysis supports the findings of Pathak et al.
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(1998) for different genotypes and environments. However, more than 100%
heterobeltiosis was observed by S. K. Dhankhar et al. (1996) and Thippeswamy
(2001) utilizing this statistic for dissimilar parents and environments.
For number of seed per fruit, the range of heterobeltiosis was −20.21% to
13.30%. Heterobeltiosis for this character has been reported by Ahmed et al.
(1999) for other genotypes and environments.
Of nine hybrids, two had significant positive heterosis for fruit ascorbic
acid content over the better parent. The range of heterobeltiosis was
−13.91% to 4.55%. The maximum heterobeltiosis occurred in the ‘Deb-401’ ×
‘Parbhani Kranti’ cross. No information is available for heterobeltiosis of this
character.
The range of heterobeltiosis for crude protein content of fruit was −16.64%
to 3.47%. The maximum heterobeltiosis occurred in the ‘Deb-401’ × ‘Bengal
IMP’ cross. Heterosis for this character has been reported by Shoba and
Mariappan (2007) for different genotypes and environments.
For fruit yield per plant, three of nine hybrids had significant positive
heterosis over the better parent. The range of heterosis for fruit yield was
−32.78% to 36.27% over the better parent. Good crosses for heterobeltiosis
were ‘VNR Green’ × ‘Shagun’ followed by ‘Barsha Laxmi’ × ‘Parbhani Kranti’
and ‘VNR Green’ × ‘Bengal IMP’. The best hybrid on the basis of mean per-
formance was ‘VNR Green’ × ‘Shagun’ with a heterosis of 36.27% over the
better parent. Heterosis for this character by Line × Tester analysis supports
the findings of Jaiprakashnarayan et al. (2008) for dissimilar parents and envi-
ronments. However, the maximum heterobeltiosis of 166.16% for this character
utilizing the present method of analysis was reported by S. K. Dhankar et al.
(1998) for different parents.
The heterosis over the better parent for PDI of YVMV ranged from
−12.95% to 140.82%. Selection of hybrids with negative heterosis over
their better parents for this character may be useful for developing YVMV-
tolerant hybrids. Good crosses for heterobeltiosis in the desired direction
were ‘VNR Green’ × ‘Shagun’, ‘Barsha Laxmi’ × ‘Parbhani Kranti’, and ‘Deb-
401’ × ‘Parbhani Kranti’. These crosses, involving Resistant × Moderately
Susceptible parents, were resistant, with very low PDI values suggest-
ing the presence of a single dominant gene controlling the resistance to
YVMV. In the crosses ‘Punjab-8’ × ‘Pusa Sawani’ and ‘Parbhani Kranti’ ×
‘Pusa Makhmali’, the presence of a single dominant gene controlling YVMV
 Breeding Okra 71

resistance was confirmed (Arora et al., 2008) along with some minor genes,
which seemed to act as modifiers in the presence of a major gene. Arumugam
and Muthukrishnan (1980) and Jambhale and Nerkar (1981) found a single
dominant gene from the cross of Abelmoschus esculentus × A. manihot and
A. esculentus × A. manihot ssp. Manihot, respectively. However, the presence
of a major gene along with minor genes for resistance to YVMV indicates
that the resistance mechanism to the virus is not as simple as reported by
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earlier workers. Resistance genes of major effect can be transferred to the

adapted varieties, but the resistance-breaking strains of the virus may not
allow resistance in these varieties to last long. Vanderplank (1984) suggested
that to achieve stable resistance, we must accumulate gene effects and contin-
uously variable resistance, which are inherited additively and have a record of
stability in the face of pathogen variability.
Analyses revealed that the crosses ‘VNR Green’ × ‘Shagun’ and ‘Barsha
Laxmi’ × ‘Parbhani Kranti’, which had maximum heterosis for fruit yield per
plant, also exhibited heterosis over better parent for plant height, number
of nodes on the main stem, number of leaves per plant, fruit diameter, fruit
weight, and number of fruits per plant. Heterosis for fruit yield per plant could
be ascribed mainly to heterosis observed for plant height, number of nodes on
the main stem, number of leaves per plant, fruit diameter, fruit weight, and
number of fruits per plant.
Specific combining ability effects represent dominance and epistatic com-
ponents of genetic variation that are not fixable, but crosses with high sca
effects involving good general combiner parents can be exploited in breeding
programs. The ‘VNR Green’ × ‘Shagun’ cross indicated significant positive sca
effects for 10 characters, namely, plant height, number of nodes on the main
stem, number of leaves per plant, fruit length, fruit diameter, fruit weight,
number of fruits per plant, number of seeds per fruit, crude protein content of
fruit, and fruit yield per plant. The ‘VNR Green’ × ‘Shagun’ cross had signifi-
cant negative sca effects for days to first flowering, nodes at first flowering, and
PDI, and it was highest for yield/plant in respect to sca combined with mean
performance. The cross ‘Barsha Laxmi’ × ‘Parbhani Kranti’ exhibited signifi-
cant positive sca effects for plant height, number of nodes on the main stem,
number of leaves per plant, fruit weight, number of fruits per plant, fruit crude
protein content, and fruit yield per plant and significant negative sca effects
for days to first flowering, nodes at first flowering, and PDI, with a very high
mean value for fruit yield per plant. The cross ‘Barsha Laxmi’ × ‘Bengal IMP’
also showed significant sca effects for 5 characters, namely, number of nodes on
the main stem, fruit weight, number of fruits per plant, ascorbic acid content of
fruit, and fruit yield per plant in the desired direction. The mean performance
of the hybrid ‘VNR Green’ × ‘Shagun’ was highest for fruit yield per plant. The
heterotic expression for fruit yield per plant in the ‘Barsha Laxmi’ × ‘Parbhani
Kranti’ cross was likely due to additive and Additive × Additive types of gene
 72 S. Das et al.

effects because the cross-combination involved parents with the best general
combining ability.
In general, the majority of heterotic crosses involving Poor × Good com-
biners indicates the role of combining ability diversity and genetic diversity in
realizing heterosis. The prerequisite for a high, uniform, and stable heterotic
effect is the correct gene content, which can be assembled in the homozygous
state, or, if the appropriate alleles are completely dominant, as a heterozy-
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gote without affecting performance (Jinks, 1983). Crosses with high mean
performances did not exhibit the maximum heterobeltiosis for plant height,
fruit length, fruit diameter, fruit weight, number of fruit per plant, and fruit
ascorbic acid and crude protein contents. This indicates that high heterosis
was not necessarily responsible for high mean performance, or vice versa,
because high heterotic response of a hybrid may be due to poor performance
of its parents. Mean performance seems to be more appropriate for selecting
the best cross-combinations compared to heterotic effects. All crosses pro-
duced negative or low positive heterobeltiosis for number of leaves per plant,
fruit diameter, fruit weight, and fruit ascorbic acid and crude protein con-
tents. High-performing parents with poor gca may not produce highly heterotic
crosses. Superior performance of hybrids for number of leaves per plant, fruit
diameter, fruit weight, and fruit ascorbic acid and crude protein contents
depends on the gca of the parents involved. Progress in improving the desired
trait will be slow if parental selection is based on mean performance alone. For
continued improvement, selection of parents should be based on mean perfor-
mance as well as combining ability and heterosis. The absence of significant
heterosis in most of the crosses could be explained by internal cancellation of
heterosis components. In the present study, genetic divergence among parental
lines is unknown and the only recourse is to determine levels of genetic diver-
gence empirically by means of variety crosses. Genetic divergence of parental
varieties is inferred from the heterotic patterns manifested in the series of
variety crosses. Heterosis in some crosses was relatively large in the present
study. Parents involving these crosses were genetically more diverse than two
other parents that manifest little or no heterosis in their crosses.
Normally, sca effects do not contribute much to improvement of crops like
okra that are often cross-pollinated. However, crosses showing desirable spe-
cific and good general combining ability could be utilized in breeding programs.
Those programs would be more effective if the parents are good combiners or
any one of them is a poor combiner. The crosses ‘Barsha Laxmi’ × ‘Parbhani
Kranti’ and ‘VNR Green’ × ‘Shagun’ were identified as good specific combin-
ers for their high sca effects for fruit yield per plant and other contributing
characters. One parent involved in these two crosses had a high gca effect and
high mean performance for several characters. Parents with High × High or
High × Poor gca effects could produce desirable transgressive segregants in
advance generations because additive genetics present in the good combiner
 Breeding Okra 73

and complementary epistatic effects in the F1 may act in the same direction to
maximize desirable plant attributes.

Disease Reaction of Parental Lines and Hybrids

Screening techniques of okra genotypes against YVMV use percentage dis-
ease incidence at different DAS. Information regarding evaluation of genotypes
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for disease reaction by disease index is lacking. Reactions of different lines and
testers and their corresponding hybrids in terms of PDI values of YVMV dif-
fered at different DAS for okra. All parents and hybrids had comparatively
lower PDI values from 30 to 60 DAS, with PDI values varying from 0.0% to
10.74% in parents and hybrids, respectively (Table 5). The PDI values among
parents were lower in ‘Deb-401’, ‘Barsha Laxmi’, and ‘VNR Green’ and higher

Table 5: Periodical observations on percentage disease index (%) of YVMV disease.

Percentage disease index

Parent/hybrid 30 DAS 45 DAS 60 DAS 75 DAS

Barsha Laxmi 0.00 0.34 2.84 8.72

(0.50)a (3.30) (9.64) (17.15)
Deb-401 0.00 0.23 2.39 7.74
(0.50) (2.74) (8.76) (16.14)
VNR Green 0.05 0.85 3.75 10.29
(1.10) (5.20) (11.09) (18.69)
Parbhani Kranti 0.12 1.10 7.65 16.68
(1.77) (5.97) (16.03) (24.10)
Bengal IMP 0.19 3.08 10.18 23.36
(2.44) (10.09) (18.60) (28.88)
Shagun 0.12 2.28 9.38 19.37
(1.78) (8.53) (17.81) (26.09)
Barsha Laxmi × Parbhani Kranti 0.01 0.50 1.91 7.72
(0.60) (3.95) (7.86) (16.13)
Barsha Laxmi × Bengal IMP 0.06 1.39 3.65 12.95
(1.32) (6.77) (11.02) (21.09)
Barsha Laxmi × Shagun 0.05 1.19 3.41 9.47
(1.20) (6.24) (10.56) (17.90)
Deb-401 × Parbhani Kranti 0.00 0.56 2.15 7.21
(0.50) (4.22) (8.35) (15.58)
Deb-401 × Bengal IMP 0.11 1.48 9.23 16.93
(1.80) (6.96) (17.68) (24.28)
Deb-401 × Shagun 0.12 2.36 10.74 18.64
(1.81) (8.83) (19.11) (25.57)
VNR Green × Parbhani Kranti 0.09 0.99 4.37 10.32
(1.51) (5.58) (12.00) (18.73)
VNR Green × Bengal IMP 0.21 2.36 9.83 17.69
(2.50) (8.76) (18.24) (24.86)
VNR Green × Shagun 0.10 1.11 3.37 9.11
(1.62) (6.03) (10.53) (17.55)
SE (±) 0.460 0.584 0.774 0.600
Least significant difference at P ≤ 0.05 1.333 1.692 2.242 1.738
a Values in parentheses indicate angular transform values.
 74 S. Das et al.

in ‘Bengal IMP’ and ‘Shagun’. The PDI values were lowest in ‘Barsha Laxmi’ ×
‘Parbhani Kranti’ and highest in ‘Deb-401’ × ‘Shagun’ crosses up to 60 DAS.
All parents exhibited more or less the same responses but hybrids had variable
PDI responses at 75 DAS.
Parental lines ‘Deb-401’ and ‘Barsha Laxmi’ were resistant; ‘VNR Green’
was moderately resistant; ‘Parbhani Kranti’, ‘Bengal IMP’, and ‘Shagun’
were moderately susceptible. Among hybrids, ‘Deb-401’ × ‘Parbhani Kranti’,
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‘Barsha Laxmi’ × ‘Parbhani Kranti’, ‘VNR Green’ × ‘Shagun’, and ‘Barsha

Laxmi’ בShagun’ had lower PDI values and are considered resistant; ‘VNR
Green’ × ‘Parbhani Kranti’ and ‘Barsha Laxmi’ × ‘Bengal IMP’ were moder-
ately resistant, and the rest were moderately susceptible.
Nonadditive gene effects in governing fruit yield attributes, quality traits,
and PDI of YVMV are important in okra. Among parental lines, ‘Barsha Laxmi’
and ‘Parbhani Kranti’ were the best general combiners for many traits and
could be used in okra hybridization programs. The results of the crosses of
‘VNR Green’ × ‘Shagun’ and ‘Barsha Laxmi’ × ‘Parbhani Kranti’ could be
exploited commercially due to the high yield coupled with low PDI values for
YVMV.

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