Diallel analyses of soluble sugar content in cowpea (Vigna unguiculata L.

Walp.)

Maina Antoine Nassourou, Tchiagam Jean Baptiste Noubissié, Yanou

Nicolas Njintang, Joseph Martin Bell

PII: S2214-5141(17)30055-7
DOI: doi:10.1016/j.cj.2017.05.005
Reference: CJ 238

To appear in: The Crop Journal

Received date: 27 December 2016

Revised date: 7 May 2017
Accepted date: 7 May 2017

Please cite this article as: Maina Antoine Nassourou, Tchiagam Jean Baptiste
Noubissié, Yanou Nicolas Njintang, Joseph Martin Bell, Diallel analyses of solu-
ble sugar content in cowpea (Vigna unguiculata L. Walp.), The Crop Journal (2017),
doi:10.1016/j.cj.2017.05.005

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 ACCEPTED MANUSCRIPT
Diallel analyses of soluble sugar content in cowpea (Vigna unguiculata L. Walp.)

Maina Antoine Nassouroua,*, Tchiagam Jean Baptiste Noubissiéa, Yanou Nicolas Njintanga,b, Joseph Martin Bellc
a
Department of Biological Sciences, Faculty of Science, University of Ngaoundéré, P.O. Box 454 Ngaoundéré, Cameroon
b
Department of Food Science and Nutrition, ENSAI, University of Ngaoundéré, P.O. Box 455 Ngaoundéré, Cameroon
c
Genetics and Plant Breeding Unit, Department of Plant Biology, Faculty of Science, University of Yaoundé I, P.O. Box 812 Yaoundé, Cameroon

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Abstract: The content of soluble sugar in cowpea (Vigna unguiculata L. Walp.) seeds is important for their
acceptability to consumers. The objective of this study was to evaluate the genetic control of soluble sugar content

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in cowpea using diallel analysis. For this purpose, half-diallel crosses were made among seven diverse parents.

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The amount of total soluble sugar was estimated by the phenol sulfuric acid reagent method and reducing sugar
were measured by the DNS (3,5-dinitrosalicylic acid) reagent method. The total soluble sugar ranged from 11.12
(VYA) to 40.79 mg g1 (IT93K-693-2) with a mean of 22.31 mg g1. Reducing soluble sugar showed a mean

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value of 9.11 mg g1 and ranged from 13.61 (TVx 3236) to 4.98 mg g1 (Lori niébé). Diallel analysis showed that
both additive and non-additive gene effects were significant in the genetic control of these traits. However,
dominance variance was greater than additive variance for soluble sugar. The traits were controlled mainly by an
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overdominance model, suggesting the advantage of delaying selection to later generations. Soluble sugar content
was found to be highly heritable, with broad-sense heritability ranging from 0.987 to 0.976. Narrow-sense
heritability ranged from 0.154 to 0.422 and revealed the lower importance of additive variance. Parents and F1
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hybrids differed significantly on the basis of their general and specific combining abilities, respectively. Recessive
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alleles had positive effects on total soluble sugar and reducing soluble sugar. These results could help cowpea
breeders improve the acceptability of cowpea seeds and cowpea-based foods by appropriate selection.
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Keywords: Cowpea; Genetic improvement; Diallel analysis; Soluble sugar

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1. Introduction

Cowpea (Vigna unguiculata L. Walp.) is an important food legume in Africa and other tropical zones. Its seeds
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are rich in proteins, carbohydrates, calories, vitamins, and essential minerals for human nutrition in many
countries [1–3]. Cowpea varieties have different tastes, based mainly on soluble sugar in the seeds; the sweet
genotype 24-125B is a favorite of Cameroonian farmers [4]. Indeed, desirable sugars such as sucrose (a non-
reducing soluble sugar), and glucose and fructose (reducing soluble sugar) can help improve the taste and flavor
of legume seeds [5, 6]. Sucrose content constituted 71% of total soluble sugar in vegetable soybean [7]. Sweetness
could help increase consumption of dry seeds and acceptance of cowpea-based food products [8].

One of the objectives of cowpea improvement is to develop cultivars with greater sucrose content that would
confer a sweet taste on the seeds [1, 8]. Very limited efforts have been made for genetic improvement of soluble
sugar in these seeds. Thus, Noubissié et al. [9] reported that sucrose content is controlled by additive and non-
additive genes and is highly heritable. Other studies of the genetics of sucrose content, conducted in soybean
(Glycine max (L.) Merr.), showed that additive and non-additive gene effects were significant [10, 11].

*
Corresponding author: Maina Antoine Nassourou, E-mail address: mainantoine@gmail.com.
Received: 2016-12-27; Revised: 2017-05-07; Accepted: 2017-06-19.

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Diallel crossing is commonly adopted to evaluate the general and specific combining abilities of parental and
hybrid lines, respectively, for traits of interest. This mating design has also been identified by Mather and Jinks
[12] as a tool for evaluating genetic components underlying the inheritance of quantitative traits. Knowledge of
heritability and combining abilities of inbred lines for soluble sugar is essential for efficient breeding of sweet
cultivars in the Sudano-Sahelian zone of Cameroon. The objectives of this study were to evaluate the contents of

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total soluble sugar and reducing soluble sugar and elucidate their genetic control in order to propose a suitable
breeding strategy for improving the sweetness of cowpea seeds.

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2. Materials and methods

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2.1. Plant materials and field experiments

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Field experiments were conducted from 2011 to 2014 at the Institute of Agricultural Research for Development
(IRAD) farm of Giring-Maroua (09°30′ N, 10°32′ E) located in the Sudano-Sahelian zone of Cameroon.

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Seeds of 15 fully homozygous cowpea cultivars (two local landraces and 13 improved lines) were obtained
from the IRAD-Maroua. The physical properties of the seed varieties were analyzed according to Baryeh [13] and
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are summarized in Table 1. Preliminary trials were conducted during the rainy season in 2011 and 2012 to ensure
purity of genotypes and to assess their variability for soluble sugar. The experiments were run in a randomized
complete block design (RCBD) with three replicates.
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Table 1 – Seed characteristics of a set of cowpea pure lines.

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Testa 100-seed Seed dimensions (cm)b

Genotype Seed color a
texture weight (g) Length Width Thickness
24-125B White Rough 16.04 ± 0.24 0.734 ± 0.006 0.615 ± 0.003 0.482 ± 0.008
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B301 Brown Smooth 9.71 ± 1.61 0.542 ± 0.013 0.502 ± 0.008 0.399 ± 0.003
Banki White Rough 18.70 ± 0.95 0.801 ± 0.027 0.495 ± 0.009 0.583 ± 0.020
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BR1 White Rough 24.27 ± 0.31 0.905 ± 0.033 0.717 ± 0.014 0.581 ± 0.002
CRSP niébé White Rough 25.06 ± 0.50 0.897 ± 0.010 0.768 ± 0.014 0.558 ± 0.001
IT93K-693-2 White Rough 21.49 ± 0.18 0.913 ± 0.012 0.647 ± 0.009 0.543 ± 0.019
IT97K-573-1-1 White Rough 22.69 ± 0.23 0.974 ± 0.017 0.666 ± 0.005 0.547 ± 0.003
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IT97K-819-118 White Rough 17.28 ± 0.87 0.776 ± 0.022 0.613 ± 0.016 0.497 ± 0.011
IT98K-205-8 White Rough 17.16 ± 0.30 0.829 ± 0.006 0.627 ± 0.008 0.470 ± 0.010
IT98K-491-4 White Rough 18.75 ± 1.13 0.893 ± 0.019 0.498 ± 0.016 0.608 ± 0.011
Lori niébé White Rough 24.68 ± 0.29 0.906 ± 0.013 0.740 ± 0.003 0.569 ± 0.006
TN-5-78 Brown Smooth 19.55 ± 0.52 0.863 ± 0.010 0.725 ± 0.010 0.561 ± 0.011
TVX 3236 Cream Rough 11.12 ± 0.46 0.625 ± 0.009 0.552 ± 0.004 0.449 ± 0.014
VYA White Rough 21.19 ± 0.26 0.825 ± 0.009 0.708 ± 0.010 0.516 ± 0.006
Warba White Rough 18.19 ± 0.60 0.849 ± 0.018 0.512 ± 0.002 0.613 ± 0.005
Mean 19.06 ± 4.45 0.822 ± 0.113 0.626 ± 0.094 0.532 ± 0.061
LSD0.05 1.36 0.035 0.029 0.019
CV (%) 23.37 13.796 14.979 11.398
a
Mean ± standard deviation of 100 measurements from triplicate determination.
b
Mean ± standard deviation of 25 measurements from triplicate experiments.
LSD, Least significant difference; CV, coefficient of variation.

Seven genotypes (24-125B, B301, BR1, CRSP niébé, IT97K-573-1-1, Lori niébé, and VYA) were selected as
parents for diallel crossing on the basis of their genetic variability for these traits. Seeds of these parents were
sown during the 2013 rainy season for crossing [14]. All 15 pure lines and the 21 F1 hybrids obtained were
planted in a RCBD with three replications during the 2014 rainy season.

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2.2. Biochemical analysis

A random sample of 0.5 g flour made from seeds of each genotype according to the procedure of Phillips et al.
[15] was used for the ethanolic extraction of soluble sugar. Seeds were extracted with 15 mL of 80% ethanol
following Meena et al. [16] and the extracts were used for all biochemical analyses.

The amount of total soluble sugar (TSS) was estimated by the phenol sulfuric acid reagent method [17]. The

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quantity of TSS was expressed as mg per g dry weight (DW).

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The content of reducing soluble sugar (RSS) was determined using the method of Miller [18]. The quantity of

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RSS was expressed as mg per g DW.

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2.3. Data analyses

Soluble sugar data were subjected to analysis of variance (ANOVA) data using STATGRAPHICS Plus 5.0 [19].

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The genetic analysis was performed for a 7 × 7 half-diallel mating population using the DIAL98 computer
program [20].

ANOVA by the method of Walters and Morton [21] was used for the study of genotypic effects partitioned into
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additive effects (a) and dominance effects (b).

Genetic parameters were estimated by Hayman’s method [22]. Heritability in the broad sense (h2) was
estimated as the ratio of total genetic variance to phenotypic variance and heritability in the narrow sense (h2n)
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was estimated as the ratio of additive variance to phenotypic variance [12]. To verify conformity to an additivity-
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dominance model, the coefficient of correlation R was calculated using Wr covariance and Vr and Vp variances (1):
𝑊r
𝑅= (1)
√𝑉r +𝑉p
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If R ≤ 1, the scatter points are within the limiting parabola 𝑊𝑟2 = 𝑉r × 𝑉p

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Griffing’s method 2 (excluding reciprocal F1 crosses) and model 1 (fixed effects) [23] were used to estimate the
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general combining ability (GCA) of the lines and the specific combining ability (SCA) of crosses. GCA and SCA
estimates of parents and hybrids, respectively, were obtained as (2) and (3):

GCAi = Xi – X (2)

SCAi = Xij – Xi – Xj + X (3)

where X is the general mean of the population, Xi is the mean of the hybrids from parent i, Xj is the mean of the
hybrids from parent j, and Xij is the value of the hybrid from parents i and j. Student’s t test was used to test the
hypotheses that the GCA or SCA effects equal zero.

Heterobeltiosis (HB%) was estimated as the deviation of the F1 value (F1) from the better parent value (BP) as
suggested by [24]. The following formula was used (4):
F1 −BP
HB% = BP
× 100 (4)

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3. Results

3.1. Genotypic variation for soluble sugar content

The ANOVA for TSS and RSS showed a significant difference (P < 0.001) between the genotypes studied (Table
2). The varieties with the greatest TSS were IT93K-693-2, BR1, and IT98K-491-4 and those with the lowest were
VYA, TN-5-78, and TVx 3236. For RSS content, accounted for mainly by simple sugars, the varieties with the

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greatest RSS were TVx 3236 and IT98K-205-8, and those with the lowest were Lori niébé and Banki.

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Table 2 – Sugar contents of 15 cowpea pure lines.
Genotype Total soluble sugar (mg g1) Reducing soluble sugar (mg g1) Non reducing soluble sugar (mg g1)

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24-125B 19.21 ± 0.12 10.65 ± 0.74 8.56 ± 0.85
B301 18.83 ± 0.07 10.57 ± 0.16 8.26 ± 0.23
Banki 19.15 ± 0.22 6.27 ± 0.61 12.88 ± 0.81
BR1 40.64 ± 0.09 8.76 ± 0.32 31.88 ± 0.40

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CRSP niébé 19.46 ± 0.06 9.40 ± 0.57 10.07 ± 0.63
IT93K-693-2 40.79 ± 0.81 6.95 ± 0.59 33.84 ± 0.89
IT97-573-1-1 20.15 ± 0.21 10.52 ± 0.32 9.63 ± 0.46
IT97K-819-118 18.70 ± 0.15 7.65 ± 0.32 11.05 ± 0.33
IT98K-205-8 19.03 ± 0.42 13.52 ± 0.28 5.51 ± 0.66
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IT98K-491-4 36.80 ± 0.50 8.20 ± 0.58 28.59 ± 0.66
Lori niébé 18.78 ± 0.11 4.98 ± 0.28 13.80 ± 0.27
TN-5-78 15.33 ± 0.30 8.45 ± 0.49 6.88 ± 0.57
TVx 3236 18.09 ± 0.26 13.61 ± 1.41 4.48 ± 1.19
VYA 11.12 ± 0.32 7.53 ± 0.48 3.58 ± 0.26
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Warba 18.61 ± 0.16 9.64 ± 0.15 8.97 ± 0.23

Mean 22.31 ± 8.95 9.11 ± 2.42 13.20 ± 9.68
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LSD0.05 2.73 0.74 1.42

CV (%) 40.12 26.51 73.35
LSD, Least significant difference; CV, coefficient of variation.
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3.2. Diallel analysis for soluble sugar content

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ANOVA by the method of Walters and Morton [21] revealed that additive effects (a) and dominance effects (b)
were significant (P < 0.01) for both TSS and RSS contents (Table 3). Dominance components (b1, b2, b3) were all
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significant (P < 0.01) except for b1 (mean dominance effect) for RSS.

Table 3 – Mean squares of total soluble sugar and reducing soluble sugar in cowpea seed for ANOVA in a 7  7 half-diallel
design.
Source of variation df Total soluble sugar Reducing soluble sugar
Block 2 0.23 0.08
a 6 247.00** 14.27**
**
b 21 9.76 24.46**
b1 1 24.14** 0.19
b2 6 162.42** 7.19**
b3 14 −56.70** 33.60**
Error 54 0.24 0.34
df, degrees of freedom; a, additive effect; b, dominance effect; b1, mean dominance effect; b2, additional dominance deviation due to
the parents; b3, residual dominance effect. * and ** significant at the 0.05 and 0.01 probability levels, respectively.

Broad- and narrow-sense heritability for TSS and RSS ranged from 0.976 to 0.987 and from 0.154 to 0.422,
respectively (Table 4). The average degree of dominance for both traits was greater than 1.0. Parents contributed
48% to 85% of dominant genes for RSS and TSS, respectively (Table 4). The direction of dominance was
negative for the two traits studied (Table 4).

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Table 4 – Estimates of ratios between genetic components in the F1 population of a half-diallel cross observed for soluble
sugar in cowpea seed.
Genetic parameter Total soluble sugar Reducing soluble sugar
Average degree of dominance (H1/D)1/2 1.10 2.19
Proportion of dominant genes 0.85 0.48
Direction of dominance (h) –2.12 −0.19
Broad-sense heritability (h2) 0.987 0.976

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Narrow-sense heritability (h2n) 0.154 0.422
Coefficient of correlation (R) 0.278 0.440

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r (Pr, Vr+Wr) 0.86* 0.67ns
Regression (Pr, Vr+Wr) 4.7Pr–74.44 1.14Pr–0.10
D, additive variance; H1, dominance variance 1; r (Pr, Wr + Vr), correlation between the degree of dominance of the parents (Wr + Vr)

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and the parental value (Pr); ns, non-significant at the 0.05 probability level; * significant at the 0.05 probability level.

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The correlations between the degree of dominance (Wr+Vr) and parental values (Pr) were positive and

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significant for TSS content (r = 0.86) but not significant for RSS (r = 0.67) (Table 4). In Fig. 1, the scatter points
are within the limiting parabola for all the traits. The correlation coefficients (R) were lower than 1 and ranged
between 0.278 (TSS) and 0.440 (RSS) (Table 4), confirming the observations of Fig. 1.
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(A) 80
 Wr = 1.161*Vr – 6.466
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P3
60

 Wr² = 82.13Vr
40

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 Wr = Vr – 4.278

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20
P7

R
P2
P5
0
Wr

P6
P4

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M P1

–20

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–40
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–60

–80
0 10 20 30 40 50 60 70 80
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Vr
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(B) 15
P

M'
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10
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 Wr² = 4.54Vr P1
5
P7  Wr = 0.451*Vr – 0.569
P4 P5
Wr

P6
 Wr = Vr – 4.307
P2
0
P3

–5

–10
0 5 10 15
Vr

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Fig. 1 – Wr/Vr graphs for total soluble sugar (A) and reducing soluble sugar (B) contents in cowpea. 𝑾𝟐𝒓 = 𝑽𝐫 × 𝑽𝐩 (limiting
parabola), where Wr is the covariance between the parents and their offspring in the rth array, Vr is the variance of the rth
array, and Vp is the variance of the parents. Red line, tangent to the limiting parabola (Wr = 1.0Vr + b); blue line, regression
of Wr on Vr. rth array corresponds to parents, P1 (24-125B), P2 (B301), P3 (BR1), P4 (CRSP niébé), P5 (IT97K-573-1-1), P6
(Lori niébé), and P7 (VYA).

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The regression line passed below the origin, and the regression coefficient of Wr on Vr varied from 0.451 (RSS)
to 1.161 (TSS) (Fig. 1). The variety BR1 corresponded to the point farthest from the origin and theoretically

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carried the most recessive alleles, whereas CRSP niébé, 24-125B, IT97K-573-1-1, Lori niébé and B301 (close to

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the origin) carried the most dominant alleles for TSS (Fig. 1). A different tendency was observed for RSS, for
which BR1 and Lori niébé had the most dominant alleles, whereas 24-125B had the most recessive alleles (Fig. 1).

The ANOVA based on Griffing’s method [23] showed that the mean squares of GCA and SCA were significant

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(P < 0.01) for both traits (Table 5). The value of σ2GCA/σ2SCA ratio showed that SCA variance was higher than
GCA variance (0.12–0.41).
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Table 5 – Mean squares for general and specific combining abilities for soluble sugar of cowpea in 7  7 half-diallel crosses.
Source of variation df Total soluble sugar Reducing soluble sugar
GCA 6 6.52** 36.13**
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SCA 14 10.74** 17.50**

Error 40 0.71 1.78
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σ2GCA/σ2SCA 0.12 0.41

df, degrees of freedom; GCA, general combining ability; SCA, specific combining ability; σ2GCA, variance of general combining
ability; σ2SCA, variance of specific combining ability; ** significant at the 0.01 probability level.
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Only 24-125B and IT97K-573-1-1 showed positive and significant GCA effects (P < 0.05) for TSS and RSS,
respectively (Table 6). In general, the studied parents showed a relatively high proportion of dominant genes for
the two traits (Table 6). The 21 hybrids showed positive or negative SCA effects (Table 7). The combinations 24-
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125B  B301, CRSP niébé  Lori niébé and IT97K-573-1-1  VYA showed positive and significant SCA effects
(P < 0.05) for TSS. For RSS content, four combinations showed a positive SCA effect: 24-125B  BR1, 24-125B
 IT97K-573-1-1, B301  VYA, BR1  Lori niébé. Positive values of SCA effects were associated with
percentage of heterobeltiosis (HB%). Indeed, for hybrids with positive and significant SCA effects, positive
percentages of heterobeltiosis were also noted (Table 7).

Table 6 – Estimates of general combining ability effects (GCA) and proportion of dominant genes (%D) for soluble sugar
among seven cowpea pure lines.
GCA effects and proportion of dominant genes (%D)
Parents Total soluble sugar Reducing soluble sugar
GCA %D GCA %D
24-125B 0.85* 38.4 1.13 17.6
B301 0.30 36.4 0.03 24.7
BR1 −0.93** 8.7 0 27.3
CRSP niébé 0.47 37.3 −1.80* 23.7
IT97K-573-1-1 −0.81* 38.2 2.30** 23.5
Lori niébé −0.10 36.8 −2.11* 27.4
VYA 0.21 32.5 0.46 23.5
Standard error 0.25 0.59
*
and ** significant at the 0.05 and 0.01 probability levels, respectively.

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Table 7 – Estimates of specific combining ability effects (SCA) and heterobeltiosis (HB%) for soluble sugar among 21 F1
cowpea hybrids.
Total soluble sugar Reducing soluble sugar
Combination
SCA effect Heterobeltiosis (%) SCA effect Heterobeltiosis (%)
24-125B  B301 3.56*** 28.04 −2.12** −29.18
24-125B  BR1 −1.51** −54.99 3.42*** 22.47
24-125B  CRSP niébé −0.72 5.27 0.19 9.98

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24-125B  IT97K-573-1-1 0.48 1.29 2.90*** 39.26
24-125B  Lori niébé −1.63** −1.07 −2.20** −49.98

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24-125B  VYA −0.18 8.07 −2.19** −25.79
B301  BR1 0.79 −50.68 −0.31 −22.31
B301  CRSP niébé −1.65** −2.34 −1.03 −46.09

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B301  IT97K-573-1-1 −0.05 −4.09 0.54 7.56
B301  Lori niébé −2.34*** −5.76 −0.25 −41.66

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B301  VYA −0.32 6.61 3.17*** 15.05
BR1  CRSP niébé 0.17 −51.80 −0.33 −27.33
BR1  IT97K-573-1-1 0.34 −54.52 −4.71*** −42.20
BR1  Lori niébé 0.23 −53.05 3.10*** 8.32

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BR1  VYA −0.01 −52.88 −1.17* −11.09
CRSP niébé  IT97K-573-1-1 −2.15*** −13.66 0.28 −11.76
CRSP niébé  Lori niébé 4.03*** 24.76 0.03 −50.75
CRSP niébé  VYA 0.32 7.32 0.85 −14.74
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IT97K-573-1-1  Lori niébé 0.45 −3.56 0.47 −12.91
IT97K-573-1-1  VYA 0.93* 0.33 0.51 11.88
Lori niébé  VYA −0.74 2.53 −1.16* −24.50
Standard error 0.35 6.05 0.44 5.61
* **
, , and *** significant at the 0.05, 0.01, and 0.001 probability levels, respectively.
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4. Discussion
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Significant differences among the tested genotypes for soluble sugar indicated large variability in the material.
Variability for soluble sugar and total carbohydrates has previously been reported for cowpea [25–27]. Total and
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reducing sugars contents reported earlier by Mune et al. [27] were relatively high compared to those obtained in
this study. These variations observed for soluble sugar content may depend on cowpea genotypes and the
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extraction methods used.

The significance of the mean dominance deviation (b1) for TSS indicates that there is a non-directional
dominance effect in the direction of the poor parent [21], and for RSS there is bidirectional dominance. The
significance of b2 (additional dominance deviation due to the parents) for these two characters suggests that the
dominant genes are not equally distributed among the parents. These latter would have an unequal number of
genes for these traits. The significance of the residual dominance effects (b3) suggests a preponderance of
dominance in the control of these traits.

The significance of the combining abilities for both traits shows the importance of both additive and dominance
effects [23]. The preponderance of SCA is shown by the values of σ2GCA/σ2SCA ratios and the variance components
for TSS and RSS content. This revealed the higher influence of non-additive gene effects [12]. Similar results
have been observed by Sadaiah et al. [28] for sugar content in sweet maize. However, this finding is contrary to
that of McPhee et al. [29] who found that estimates of GCA were on average 16.5 times higher than those of SCA
for raffinose family of oligosaccharides (RFO), suggesting that additive genetic variance was most important in
common bean. In our study, a high proportion of dominant genes suggest a preponderance of non-additive genes

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for soluble sugar. Moreover, the average degree of dominance (H1/D)1/2 was greater than 1, suggesting
overdominance for all traits.

Broad-sense heritability values were high (97.6%–98.7%) suggesting that these traits are controlled mainly by
genetic factors and are amenable to genetic improvement. Thus, it may be possible to increase levels of soluble
sugar in cowpea through genetic improvement. The low values of narrow-sense heritability (15.4%–42.2%)

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showed a predominance of non-additive gene effects. Also, a study of McPhee et al. [29] on RFOs in common

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beans revealed that narrow-sense heritability was moderate, implying that manipulation of RFO accumulation
would probably need to be done on a progeny row basis with replications. Still, it would be difficult to adopt a

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pedigree method to improve these characters when non-additive gene action predominates [12]. Accordingly, a

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suitable and effective approach could be the exploitation of heterosis in breeding hybrid cultivars.

According to Fig. 1, genetics for soluble sugar contents follows an additive–dominance model [20]. The

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coefficients of regression of Wr on Vr were not significantly different for unity in TSS (1.161). This result
indicates the adequacy of the simple additive–dominance genetic model [22]. Indeed, F1 data from several crosses
indicated that sweetness is completely recessive [30]. The association of high soluble sugar content with recessive
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alleles might present some difficulties for selection during the early generations.

However, the coefficients were significantly different from unity for RSS (0.451), suggesting the possibility of
non-allelic interaction. Significant epistasis was observed in most crosses for sucrose content in cowpea [9]. In
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sweet field corn, TSS was influenced mainly by non-additive gene effects and duplicate epistasis was also
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reported [31]. Furthermore, regression analysis in the present study showed an overdominance type of gene action
for all characters (Fig. 1). Thus, selection based on these traits could be applied in later generations. Hybrid
genotypes are likely to produce higher soluble sugar contents.
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The parental lines 24-125B and IT97K-573-1-1 showed a positive contribution to crosses that could be
exploited to develop hybrids with sweet-tasting seeds. SCA performance might be considered as a criterion for
selecting the best crosses. Combinations with high or moderate and positive GCA showing high SCA effects may
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produce transgressive forms by combining two complementary parents to increase soluble sugar contents in
cowpea seeds. These findings suggest gene dispersion and genetic interaction between favorable alleles from both
parents [24, 32].

A moderate amplitude of HB% was recorded in some combinations, in particular 24-125B × B301 and CRSP
niébé  Lori niébé for total soluble sugar, and 24-125B × BR1, 24-125B × IT97K-573-1-1, and B301  VYA for
reducing soluble sugar. Superiority of hybrids over the better parent indicates the complementary parents that
produce the most important transgressive forms [24]. Heterobeltiosis may be linked to the cumulative action of
favorable dominant genes dispersed in two parents and/or to favorable intra- or interlocus interactions, called
overdominance [12]. The association between positive SCA and HB% suggests that HB% could contribute
reliable information to the breeding procedure.

Attention should be focused on developing F1 hybrids with positive SCA derived from parents with positive
GCA to improve sweetness of cowpea seeds. Traits conditioning sweetness appear to be controlled primarily
through an overdominance model of gene action, suggesting that selection in later generations will be most
effective.

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Acknowledgments

This work was supported in part by funds from International Foundation for Science (C/5262-1). The authors are
grateful to Ukai Yasuo, University of Tokyo, Japan for providing the microcomputer program DIAL98. They are
also thankful to the Institute of Agricultural Research for Development (IRAD) of Maroua-Cameroon for kindly
providing the seed samples and the field for experiments.

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