Euphytica 96: 207�213, 1997.

207
c1997 Kluwer Academic Publishers. Printed in the Netherlands.

Genetic analysis of yield and its components in vegetable cowpea (Vigna

unguiculata L. Walp)

Pathmanathan Umaharan*, Rasiah P. Ariyanayagam & Syed Q. Haque

Department of Plant Science, The University of the West Indies, St. Augustine,
Republic of Trinidad and Tobago;
(* author for correspondence)

Received 23 April 1996; accepted 31 December 1996

Key words: clusters, inheritance, pod weight, selection, yield components, Vigna
unguiculata, vegetable cowpea,
genetics

Summary

The F2 and backcrosses of a cross between two vegetable cowpea (Vigna unguiculata
L. Walp) varieties with
contrastingcharacteristicswereevaluatedforpodyieldanditscomponents,withtheaimofunde
rstandingthegenetic
basisofthesecharacteristics.Afour-
parametermodelincorporatingtheadditive,dominanceandadditive #additive
genetic components fitted the data for pod yield and clusters per plant. The
additive and additive #additive effects
were positive and were larger than the dominance component. The relatively large
additive and the predominantly
positive dominant effects suggest that selection would be effective. Pod weight had
high broad (84%) and narrow
sense heritability (75%) and can be effectively selected for in the early
generations. The study suggested that
vegetable cowpea improvement programs should focus on selecting for clusters per
plant and average pod weight in
the early generations, while selection for dry pod yield could be delayed to later
generations. It was concluded that
pods per plant may be a useful selection criterion in multi-location trials aimed
at selecting for stability of yield.

Introduction Yield in vegetable cowpea is usually defined as

green pod yield, expressed in kg per ha. Brathwaite

Vegetable cowpea refers to varieties of cowpea (Vigna
(1982),however,reportedthatyieldexpressedasnumunguiculata L. Walp) grown for their
immature succu-ber of marketable pods per ha is of particular interest to
lent pods, popularly known as long bean, bodi, bora, producers, since vegetable
cowpea is sold by the bunsitao, snapea, snake pea and aspargus bean in different
dles in the Trinidad markets. Yield in vegetable cow-
parts of the world. The indigenous varieties of veg-pea, further, can be thought of
as a function of compoetable cowpea were climbing types. In recent times, nents
such as the number of clusters per plant (Singh &
however, many erect bushy varieties have been devel-Mehndiratta, 1970; Khan &
Stoffella, 1985), average
oped (Acosta & Petrache, 1960; Redden, 1981; Mital number of pods per cluster
(Fernandez & Miller, 1985)
et al., 1980), with the objective of increasing yields. and average pod weight (Mak
& Yap, 1977; Aggarwal

Much work has been done towards understanding et al., 1982; Brathwaite, 1982).
the inheritance of yield and yield components in grain This study seeks to
understand the genetic basis of
cowpea (Kheradnam & Niknejad, 1974; Aryeetey & yield and its components in
vegetable cowpea with the
Liang, 1973; Lal et al., 1976; Zaveri et al., 1983). In aim of developing a
strategy for improvement of yield
contrast, however, the genetics of yield in vegetable in vegetable cowpea.
cowpea has not been investigated with the exception
of reports by Mak & Yap (1977, 1980), who worked
with climbing types of vegetable cowpea. Moreover,
genetic studies are lacking with respect to components
of yield in vegetable cowpea.
Materials and methods

The parents (P1, P2), F1, F2, BC1 and BC2 of the cross
IT-81D-1228-14 #UCR193formedthegeneticmaterialforthisstudy.IT-81D-1228-
14wasdevelopedatthe
International Institute of Tropical Agriculture, while
UCR 193 was developed at University of California
Riverside. These varieties were selected from a yield
trial of 23 dwarf vegetable cowpea varieties (Umaharan, 1990), in which they had
exhibited contrasting
values for the various yield components investigated
in this study. Moreover, both varieties showed almost
identical flowering (UCR 193 flowers in 38 days while
IT-81D-1228-14 flowers in 42 days) and maturity periods (around 100 days), which
would allow the study of
yield components without the influence of these complications.

The initial crosses as well as the F2 and

backcrosspopulationsweregeneratedinaninsectprotected
greenhouse.Approximately,20F1plantswereallowed
to self pollinate to generate approximately 1500 F2
seeds. Approximately, 160�180 F1 seeds and around
250 seeds for each of the backcross generations were
generated.

A field experiment was conducted at the University

Field Station at Valsayn, to evaluate P1, P2, F1, F2,
BC1 and BC2 for yield and the various yield components. The experiment was arranged
in a randomized
complete block design with four replications. The parents, F1, and backcross
generations were allocated to
single plots per block, while the F2 population was
distributed over three plots per block. The plots for P1,
P2 and F1 consisted of three rows (2.4 m long), spaced
60 cm apart, with each row consisting of 11 planting
points. The plots for the backcross generations consisted of four rows of similar
size and those for the
F2 consisted of 6 rows. Hence, in total, the F2 was
represented by 18 rows (198 plants) per block.

The field was ploughed and later harrowed to a

fine tilth. Sixty kg ha..1 of triple super phosphate was
incorporated into the plot during the last harrowing.
Plots were established in October, 1989 and manually weeded at 25 and 40 days after
planting (DAP).
The water requirement was met entirely by rainfall.
Insecticides (Decamethrin 2.5% w/v) and fungicides
(Benomyl 50%) were sprayed at weekly interval, to
control pests and diseases. The plants were thinned to
one per planting point, during the first weeding, 25
DAP.

Data were collected from nine competitive plants

per plot for the parents and F1s. Similarly, 30 plants

were sampled per plot (90 plants per block) for the F2
and 18 plants per plot for the backcross populations.
Data were collected on number of pods per plant, dry
pod yield per plant, average pod weight, number of
productive clusters per plant and average number of
pods per cluster. Pods were harvested five times over a
period of eight weeks. Data on productive clusters per
plant were obtained during the third harvest. Average
pod weight and average number of pods per cluster
were calculated from records of pod weight per plant,
number of pods per plant and number of clusters per
plant.

Plot means were calculated for each measurement

and analyses of variance conducted to test the significance of differences between
family means. A generation mean analysis was performed on the data in
accordance to the procedure outlined by Mather &
Jinks (1971). The data were tested for the adequacy of
the additive-dominance model using the ABC Scaling
Test (Mather, 1949) and the Joint Scaling test (Cavalli, 1952), incorpporating the
weighted least square
method of Hayman (1960). The latter approach also
provided best fit estimates for the parameters in the
model. When the additive-dominance model fitted the
data, a generation variance analysis was performed
based on the method described by Allard (1960). This
provided estimates of additive and dominance components of variance, degree of
dominance, F and heritability both in broad and narrow sense.

Whentheadditive-dominancemodelfailedtofitthe
data, a six parameter model incorporating mid parental
value [m], additive effect [d], dominance effect [h] and
the three digenic interaction components, additive
.
additive [i], dominant #dominant [j] and additive
.
dominant [I] were fitted to the data, using the method
of Mather & Jinks (1971). When one or two of the
interaction components were not significant, they were
omitted from the genetic model and a four or five parameter model was fitted using
the weighted least square
method. This not only provided improved estimates of
the parameters estimated in the model but also provided a test for the adequacy of
the model based on one
or two degrees of freedom (Mather & Jinks, 1971).
Furthermore, when the additive-dominance model did
not fit the data, a generation variance analysis was not
performed and only broad sense heritability estimates
were obtained.
Table 1. Means and standard errors for pod yield and its components in P1, P2, F1,
F2, BC1
and BC2 of the cross UCR 193 #IT-81D-1228-14 in vegetable cowpea

Generation Yield yield components

Pods per plant Dry pod Clusters per Pods per Mean pod
yield, g. plant cluster fresh wgt

P1 21.4 #0.2 32.5 #0.6 11.1 #0.4 1.94 #0.06 8.1 #0.2
P2 29.6 #1.3 57.3 #1.9 19.2 #0.8 1.53 #0.01 12.4 #0.3
F1 24.6 #0.8 53.3 #0.6 13.4 #0.3 1.84 #0.03 12.9 #0.1
F2 23.6 #0.7 45.4 #1.9 12.9 #0.2 1.83 #0.02 11.2 #0.2
BC1 22.7 #0.7 40.1 #1.3 12.2 #0.5 1.83 #0.03 10.7 #0.3
BC2 25.5 #1.0 50.9 #1.6 14.8 #0.6 1.72 #0.01 12.8 #0.3

h2 (bs) 64.0 75.6 63.9 75.0 84.4

h2 (bs) refers to heritability in the broad sense. Means of the parents (P1, P2),
and F1 were based
on 36 plants, those of F2 were based on 360 plants and those of the back cross
generations (BC1
and BC2) were based on 72 plants.

Table 2. Scaling tests and estimates of genetic components of the various genetic
models for yield and yield components in the cross of UCR
193 #IT-81D-1228-14 in vegetable cowpea

Scaling test Dry pod yield Pods per plant Clusters per plant Pods per Mean pod
genetic effects cluster fresh weight
1. Joint Scaling test 3-Parameter 4-Parameter 3-Parameter 3-Parameter 4-Parameter
3-Parameter 3-Parameter
(Mather & Jinks, 1971) model model model model model model model
� m 43.2 #0.9 3.9 #2.8 24.9 #0.6 13.9 #0.34 12.5 #0.55 1.72 #0.02 10.2 #0.14
Additive (A) d 11.0 #0.9 12.0 #0.9 3.5 #0.6 3.1 #0.39 3.6 #0.42 -0.18 #0.02 2.2
#0.15
Dominant (D) h 9.9 #1.1 19.8 #3.1 -1.2 #0.9 -0.9 #0.52 0.9 #0.75 0.16 #0.04 2.7
#0.20
A #A i � 10.6 #3.1 � � 2.4 #0.72 � �
A #D j � � � � � � �
D #D l � � � � � � �
X2 (3) 13.3##
.
� 3.4ns
13.0##
.
� 6.2##
.
2.0ns
X2 (2) � 1.8ns
� � 2.4ns
� �
2. ABC Scaling Test (Mather, 1949)
Scale A -6.2* + 2.7 � -1.8 #1.5 -0.02 #1.6 � -0.12 #0.10 0.33 #0.73
Scale B -9.2*+�3.7 � -3.1 #2.5 -2.96* #1.5 � 0.06 #0.05 0.23 #0.62
Scale C -16.0* + 7.9 � -5.6 #5.6 -5.30* #1.6 � 0.15 #0.13 1.01 #1.40

######
, , refer to significance at P < 0.01 and P < 0.001, respectively.

Results

Dry pod yield

Analysis of variance of dry pod yield showed significant differences between the
parents and among
generation means (Table 1). An additive-dominance
model could not explain the variation among generation means. Nevertheless, a four
parameter model
consisting of [m] [d] [h] and [i] components fitted the
data (Table 2). The dominance component was positive

which suggested that dominance was in the direction

of more pod yield. The additive [d] and additive
.
additive [i] components were in the same direction,
reinforcing each other, and the combined [d] + [i] component was larger than the
dominant component. The
results indicate that response to selection would be
good. The broad sense heritability of 75.6, indicates
that the ambiguity of environmental influences may be
low under the experimental conditions even for yield.
Table 3. Estimates of genetic components of variance, average degree of dominance,
heritability and the ratio of F/pDH for some components of yield in Vigna
unguiculata L. Walp

Estimates Pods per plant Mean pod fresh weight Pods per cluster

Additive variance D 72.24 14.46 0.31

Dominant variance H 49.14 5.00 0.22
Error variance E 26.63 1.09 0.07

1. pH/D 0.8 0.6 0.8

2. h2 (ns) 46.1 75.6 54.8
3. h2 (bs) 64.0 84.4 73.7
4. F/pDH -0.2 0.1 -0.4
1 = degree of dominance; 2 = heritability in the narrow sense; 3 = heritability in
the broad
sense; 4 = direction and the relative constancy of H:D over all loci.

Pods per plant

Pods per plant varied from 21 in UCR 193 to 30 in

IT-81D-1228-14 (Table 1). Generation mean analysis
showed that an additive-dominance model fitted the
data. The dominant component was negative indicating that dominance was in the
direction of fewer pods
per plant and hence the F1 and F2 means were lower than the mid parent. Generation
variance analysis
showed that the additive variance was larger than the
dominance variance (Table 3). Incomplete dominance
was indicated by the data with a degree of dominance
of 0.8. The low and negative F/pDH component indicated that the dominance was in
the direction of fewer pods per plant and varied to a large extent among
the genetic determinants. The broad sense heritability
estimate was moderate (0.64), and the narrow sense
heritability was estimated to be 0.46.

Clusters per plant

Clusters per plant showed a wide variation among the

parents, with UCR 193 and IT-81D-1228-14 having

11.0 and 19.2 clusters, respectively (Table 1). The

additive-dominance model did not fit the data, but a
four parameter model incorporating [m] [d] and [i]
components fitted the data (Table 2). Dominance was
positive and was in the direction of more clusters per
plant.Theadditiveandadditive #additivecomponents
together were positive, large and reinforcing. The heritability (broad sense) was
moderately high and was
similar in magnitude (0.64) to that of number of pods
per plant (Table 1). Clusters per plant are easy to count
in the field and the study shows that its genetic nature
would allow considerable progress.
Pods per cluster

Thenumberofpodsperclustervariedfrom1.53to1.94
among the parents (Table 1). The number of pods per
cluster fitted an additive dominance model as indicated
by the non-significance of both the ABC Scaling Tests
and the Joint Scaling Test (Table 2). The estimated
genetic components from generation means indicated
the involvement of both additive and dominant effects.
The additive effect was negative indicating predominance of decreasing alleles. The
dominant effect was
positive indicating that the dominance was predominantly in the direction of more
pods per cluster. The
negative but moderately large F/pDH estimate (-0.4),
obtained in the generation variance analysis (Table 3),
confirms that dominance varies to some extent among
genetic determinants, but is largely negative. Consequently, the F1 and F2 means
were larger than the mid
parental value and a mid parental heterosis of 6% was
observed.Theestimateofthedegreeofdominancewas
0.8, which shows the relative importance of dominance
in the inheritance of this trait. The additive variation
accounted for about 50% of the total genetic variation.
Heritability in the broad sense was high (0.74).

Average pod weight

Average pod weight varied between 8.1 and 12.4 g

among the parents (Table 1). Pod fresh weight fitted
an additive dominance model (Table 2). The estimated additive and dominant
components using the generation mean analysis (Table 3) indicated that both
additive and dominant components were significant.
However, the generation variance analysis indicated
that additive variance was three times larger than dom
inance variance. The estimated degree of dominance
wa 0.6. The small positive value for F/pDH suggests
that dominance relationship may vary among genetic
determinants, but is slightly in favour of larger pod
weights. Estimates of heritability both in the broad
sense and in the narrow sense were large, indicating
that pod weight is a relatively easy trait to improve.
Further, early generation selection would be effective
for this trait. The F2 segregants varied from 5.1 to

17.3 g for pod weight and showed a large degree of

transgressive segregation, in both directions. The estimated genetic advance at 5%
selection intensity was
5.2 g.
Discussion

The large degree of variation seen among the parents

used provided an excellent opportunity to observe the
genetic basis of such differences. Although both varieties were dwarf in stature,
UCR 193 had relatively shorter and thinner pods than IT-81D-1228-14 and
consequently smaller pod weights. Further, UCR 193
had a propensity to produce more pods per peduncle,
although it had a relatively lower number of productive peduncles per plant. The
two varieties and their
progeny flowered fairly uniformly over a period of
10 days and showed fairly uniform crop senescence.
These conditions allowed the assessment of yields and
their components, without the confounding effects of
crop duration.

Greenpodyieldisgenerallyobtainedbyharvesting
pods at the green mature stage twice a week for over
a period of 6�8 weeks (Brathwaite, 1982). Because of
the intensity of work involved in harvesting individual
plants during the experiment, harvesting was done at
weekly intervals for the first three weeks and at fortnightly intervals thereafter.
This would have resulted
in the underestimation of the total yield obtained but
would not have seriously altered the relative potentials
of the plants.

Genetic analysis of yield showed that although pod

number per plant can be explained by an additive dominance model, the pod dry
weight required the incorporation of the additive #additive interaction. Although
both additive and dominant components were significant, the additive effects were
larger. The broad sense
heritability for dry pod yield was much higher (0.76)
as compared to pods per plant (0.64). This indicates
that selection for yield based on the number of pods
per plant would be less effective than that based on

pod dry weights. Bapna et al. (1972) and Fernandez &

Miller (1985), working with grain types, also observed
that pod number is the yield component most affected
by the environment. Subsequently, Ranganatha (1986)
suggested that selection for consistently high number
of pods per plant over environments is a good index of
yield stability. Mak & Yap (1980) working on climbing vegetable types also found
that non-additive gene
action was important in the inheritance of both number
of pods per plant and pod yield.

The study indicates that the combined [d] and [i]

componentsforpodyieldwaslargerthanthedominant
effects and indicated that selection would be effective
in early generations. The large and positive dominant
componentprovidestheopportunitytofurtherimprove
yields by pedigree selection within the families. However, owing to the cost of
selection for yield in the
very large early generations, selection may be strategically delayed. In the
subsequent multilocation yield
trials pod number may be used as an index of yield
stability, especially in countries where vegetable cowpea is marketed in bundles
(Brathwaite, 1982) rather
than on a weight basis. A negative correlation (-0.75)
between pod number per plant and pod length has been
reported in vegetable cowpea (Aggarwal et al., 1982)
and among grain types (Singh & Mehndiratta, 1970;
Bapna et al., 1972; Kheradnam & Niknejad, 1974).
In many countries pod length is an important criterion
governing consumer acceptability of bodi beans and
as such this may be another reason to abandon pod
number as a selection criterion in vegetable cowpea
breeding programs.

Many authors, working with grain types, have suggested that increased number of pod
bearing peduncles
(clusters) per plant may be the appropriate objective to
increase yield in cowpea (Jindla & Gupta, 1984; Khan
&Stoffella,1985).ThiswassupportedbySummerfield
et al. (1985), who concluded from a review of physiological studies in cowpea that
the number of reproductivenodesislikelytobemoreimportantthanreproductive efficiency
in explaining variations in yield. However, reported broad-sense heritabilities
were moderate(Fery,1985)andnon-additivegeneactionwasoften
found to be important in the inheritance of the trait (Lal
et al., 1976; Zaveri et al., 1983). Genetic studies in vegetable cowpea, however,
are non-existent. The present
study showed that although non-additive effects were
important, they were largely of the additive #additive
type. Moreover, the genetic effects were
predominantlyofthe[d]and[i]kindindicatingthatselectionwould
be effective.
Thenumberofpodsperpedunclehasnotbeenconsidered an important yield determinant in
grain cowpea. Lush & Evans (1981) reported that the genetic variability is limited
to 2�3 pods per peduncle
in domesticated varieties of cowpea. Littleton et al.
(1981) reported that the very high rate of pod growth
in cowpea may account for the rapid decline in photosynthetic rate of leaves.
Peoples et al. (1983) attributed
the decline in photosynthetic efficiency to a decline in
nitrogen in blossom leaves. Fruit induced senescence
is known to occur in cowpea bearing 4�6 sequentially
filling pods per peduncle (Summerfield et al., 1985).

In vegetable cowpea, the indeterminate peduncle

produces many pods sequentially without resulting in
senescence and hence pods per cluster may be a more
important determinant of yield. This may be because
the pods are harvested in a relatively immature stage in
vegetable cowpea, when the seeds are small. Further,
since the seed carries 96% of the nitrogen in the fruit
(Pate & Minchin, 1980) the demand on the nitrogen
reserves may also be lower. Chandrappa et al. (1974)
reported a high heritability for the trait, but the genetic
nature has not been investigated. This study indicated
that the additive dominance model, containing equally
large components for additive and dominance variance
explained the data. The dominance was predominantly
in the direction of higher number of pods per cluster
indicating that selection would be effective. The
negativeadditiveeffectmaylimitprogress.Thebroadsense
heritability was high (0.74).

Although, average pod weight is an important component of yield in vegetable

cowpea, the genetic nature
has not been studied. Further, pod fresh weight is
an indicator of fleshiness and pod length in vegetable
cowpea, both of which are important determinants of
market acceptability. Genetic analysis showed that an
additive dominance model explained the genetic variation in this trait. Although
both additive and dominance
componentsweresignificant,theadditivevariancewas
three times greater than the dominance variance and
consequently the narrow sense heritability was high
(84.4). The small F/pDH component (0.1) indicated
that dominance component was ambidirectional. Nevertheless, the large degree of
transgressive segregation
observed (5 g to 17 g) indicated ample opportunity for
improvement of this trait.

In conclusion, the study indicates that vegetable

cowpea improvement programs should concentrate on
selecting for clusters per plant, pods per cluster and
average pod weight in the early generations, while
selection efforts for dry pod yield may be delayed

to later generations to reduce harvesting cost. The

elite lines selected may be investigated in
multiseason/multilocationtrialswherepodnumbermaybeused
to study the relative stability/adaptabilty of the selected
lines.

Acknowledgements

The authors wish to acknowledge the technical assistance of Ms. V. Gajadharsingh

during the period of
hybridization. The financial assistance provided to the
senior author by the Government of the Republic of
Trinidad and Tobago, under the Commonwealth Fellowship Plan is gratefully
acknowledged.

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