Genetic analysis of yield and its components in vegetable cowpea (Vigna unguiculata L. Walp)

The F2 and backcrosses of a cross between two vegetable cowpea (Vigna unguiculata L. Walp) varieties with contrasting characteristics were evaluated for pod yield and its components, with the aim of understanding the genetic basis of these characteristics. A four-parameter model incorporating the additive, dominance and additive × 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. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of seed size in cowpea (Vigna unguiculata (L.) Walp.)

Summary The inheritance of seed weight in cowpea was examined in a field planting of the parents, reciprocal F₁s, F₂s and backcrosses to both parents of a cross between TVu 1977-OD (small seeded) and ACC 70002 (large).Seed weight was inherited quantitatively and small seed was partially domiminant to large seed size. Gene action was predominantly additive but dominance and additive × additive epistatic effects were also significant.Broad and narrow sense heritabilities were 85.1±5.3% and 75.4±18.6% respectively. The minimum number of loci involved in the inheritance of seed size was eight, and each gene pair contributed up to 1.02 g increase to seed weight. The estimate of genetic advance from F₂ to F₃ generations with 5% selection intensity was 3.58 g.International Institute of Tropical Agriculture, Semi-Arid Food Grain Research and Development Project/National Cowpea Improvement Program, B.P. 1783, Ouagadougou, Upper Volta.Department of Primary Industries, Hermitage Research Station, Warwick 4370, Queensland, Australia.International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), ICRISAT Center, Patancheru, P.O. 502 324, A.P., India.     

Adaptation and competition in mixtures of cowpea (Vigna unguiculata (L.) Walp.)

Summary Six pure lines and four mixtures were grown in six environments throughout lowland Papua New Guinea. There were no significant transgressive increases in the grain yield of mixtures above their pure line components, and mixture yields were adequately predicted by the mean of components. In yield stability it was found that the individual buffering of pure lines was of more importance than population buffering, and that the magnitude of population buffering varied with the particular combination of components. Competitive effects in all the mixtures were of the compensating type. Dramatic changes in mixture composition resulting from natural selection precluded their use in local agriculture. The outcome of competition in mixtures was strongly influenced by the growing environment, such that selective index was correlated to general fertility.     

Resistance to cowpea severe mosaic virus, determined by three dosage dependent genes in Vigna unguiculata L. Walp

P1, P2, F1, F2, BC1 and BC2 generations of four resistant × susceptible crosses and three resistant × resistant crosses of cowpea (Vigna unguiculata L. Walp) were screened for resistance to cowpea severe mosaic virus (CPSMV), in an insect protected screen house. The segregation ratio, at maturity, showed a 63 susceptible : 1 resistant ratio in the F2 generation indicating that resistance is governed by three major genes. The backcross tests and the F3 test confirmed this. The intermediate levels of symptoms observed in the F1 generation and the progression of symptom expression in the F2 generation suggested that resistance is gene dosage dependent. The genetic basis of immunity, tolerance, high level of resistance is discussed based on the three gene model. Delayed symptom expression was evident in genotypes with one or two susceptible alleles. The implications of delayed expression of symptoms in selection and breeding for resistance to CPSMV are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of Eye Pattern and Seed Coat Colour in Cowpea (Vigna unguiculata [L.] Walp.)

The F₂ progenies of crosses between several cowpea (V. unguiculata) lines were investigated for variation of eye pattern and seed coat colour. It was found that three (W, H, O) and five (R, P, B, M, N) major genes control eye pattern and seed coat colour, respectively. The recessive gene (GO) for restricted eye pattern enables the underlying basic white or cream seed coat colour to be observed. A similar effect is obtained with the recessive gene (rr) for colour expression. The expression of mottling (V), possibly a seed coat pattern, may for be observed when it is combined with the genes for certain eye patterns. The significance of these findings in breeding for consumer preference for specific seed coat colour is discussed.     

Linkage relationships among loci controlling morphological traits in cowpea (Vigna unguiculata (L.) Walp.)

Summary Linkage among loci controlling various morphological traits in cowpea were determined using F₂ progenies. Data were collected on individual plants of four crosses segregating for several loci. Recombination estimates between the following pairs of loci were as follows: Sw (swollen vs normal stem base)-Fbc (cream vs green flower buds) (41±4.8%), Pu^s(purple vs green stems)-Cbr (cocoa-brown vs straw-yellow dry pods) (31±5.7%), Pu^p(purple vs green immature pods)-Cbr (30±5.7%), Pu^s-Pu^p (4±1.5%), Ndt (non-determinate vs determinate)-Pd (peduncle colour) (26±2.8%), Ndt-Hg (semi-erect vs erect plant type) (26±2.8%), P^t(purple vs green pod tips)-Bk (greyish-black vs straw-yellow dry pods) (19±2.4%) and Hg-Bpd (normal vs branching peduncle (24±9.5%). Four linkage groups (LG) were identified in these studies. The proposed LG I contained loci Sw and Fbc; LG II loci Pu^s, Pu^p, and Cbr; LG III loci Pd, Ndt, Gh, and Bpd; and LG IV loci P^tand Bk.     

Evaluation of parent selection methods for yield improvement of cowpea (Vigna unguiculata (L.) Walp.)

Summary The mean aim of this study was to identify an effective method of predicting cross-potential in respect of yield improvement of the cowpea (Vigna unguiculata (L.) Walp.) through hybridization and subsequent selection.Three prediction tests based on data collected from experiments with ten cowpea lines and the F1 and F2 generations of all possible crosses between these lines were evaluated. The three prediction tests were based on: i Yields of prospective parents. ii. General combining abilities estimated from F1 and parental data iii. Frequencies of superior plants in F2 populationsEach of the three procedures was used to identify the crosses with the lowest or highest yield potential, i.e. those crosses most likely to produce the greatest number of lines with yields higher or lower than those of either parent. Crosses selected on the basis of these criteria were evaluated in the following growing season on the basis of biometrical analysis of F2 and backcross populations and the performance of the respective F3 and F4 generations.Six different crosses were selected from the 45 tested on the basis of the three prediction tests. Analyses of F2 and backeross populations from each cross provided estimates of [d] the difference in parental means and 1/2D the additive genetic component which, in turn, were used to predict the frequency of transgressive inbred lines obtained by single-seed descent from the F2. Both crosses selected on the basis of F2 data were expected to produce a much higher frequency of transgressive lines than those selected on the basis of parental yields. The high potential cross selected on the basis of general combining ability was expected to produce a higher frequency of transgressive lines than the corresponding low yield potential cross. Replicated trails of F3 bulks of the selected crosses showed that high yield potential crosses were significantly higher yielding than low yield potential crosses for all prediction criteria. In a trial with 22F4 lines of each of the two crosses selected on the basis of parental yields and frequency of high yielding F2 plants, the high yield potential cross in each case had a greater mean and variance for seed yield than the corresponding low yield potential cross.It was concluded that parental yields provide a sound basis for an initial screening of prospective parents. This screening should identify a high yielding line which can be crossed with a number of contrasting lines to produce F2 populations for the final screening phase.     

Inheritance of resistance to Striga gesnerioides biotypes from Mali and Niger in cowpea (Vigna unguiculata (L.) Walp.)

Pot and laboratory experiments were performed in order to elucidate the genetics of resistance of three cowpea (Vigna unguiculata (L.) Walp.) resistant sources to two biotypes of the parasitic weed Striga gesnerioides (Willd.) Vatke (witchweed). B301, Suvita-2 and IT82D-849 were crossed to susceptible cultivars. Roots of each plant of parental, F₁, F₂, BC₁ and BC₂ populations were washed free of soil and examined closely for S. gesnerioides attachment. Data on resistant and susceptible cowpea plants were analyzed using the chi-square test to ascertain the goodness of fit to different genetic ratios. Segregation patterns indicate monogenic dominant inheritance of resistance to both S. gesnerioides biotypes in B301. Suvita-2, which is susceptible to the biotype from Maradi, Niger, showed segregation patterns that indicate monogenic dominant inheritance of resistance to S. gesnerioides biotype from Cinzana, Mali. Results also indicate monogenic dominant inheritance of resistance to S. gesnerioides biotype from Cinzana in IT82D-849. However, the inheritance of resistance to the biotype from Maradi would be monogenic and recessive in this line. These results suggest that the resistance of all three sources would be easy to transfer to promising breeding lines. This revised version was published online in July 2006 with corrections to the Cover Date.     

Segregation Pattern of Decumbent, Climbing and Bushy Growth Habits in Vigna unguiculata (L.) Walp.

F₁ and F₂ data from crosses of cowpea accessions with three different growth forms showed that the inheritance of growth habit is governed by two allelic pairs, AA and BB. The genetic ratio obtained implicated a modifier gene interaction in which the genotype A-B- has the decumbent growth habit; the genotype aaB- has the climbing and aabb has the bushy habit. While the decumbent growth habit was dominant and epistatic over the climbing habit, the decumbent and the climbing growth forms were each monogenically dominant over the bushy habit.     

The domestication and improvement of cowpeas (Vigna unguiculata (L.) Walp.)

Summary Physiological and morphological characteristics of the two wild and three domesticated subspecies of cowpeas are compared. The wild accessions are alike in having small, hard seeds borne in dehiscent pods, but differ in other characteristics. We suggest that the wild subsp. dekindtiana, from the seasonally-arid tropics, is more likely to have been the progenitor of modern cowpeas than the other wild subspecies (subsp. mensensis), but that subsp. dekindtiana was first cultivated in the humid tropics where its pods are slow to dehisce. Domestication has been associated with changes in the structure of pod valves and seed coats which reduce pod dehiscence and seed hardness. Pods and seeds have increased in size, mainly by increases in the rate of dry weight accumulation, and their increase has been only partly paralleled by increase in the area of subtending leaves. There has been no increase in the maximum photosynthetic rate of leaves, but the duration of their photosynthetic activity has increased. Domesticates are less sensitive than are wild plants to some environmental controls, such as in the response of germination to temperature, but in their flowering responses to daylength both wild and cultivated forms retain sensitivity under conditions where this is of adaptive value.     

RFLP mapping of an aphid resistance gene in cowpea (Vigna unguiculata L. Walp)

Summary Restriction fragment length polymorphism (RFLP) analysis has several advantages over traditional methods of genetic linkage mapping, one of these being the starting point for map-based cloning. The recent development of an RFLP map of cowpea (Vigna unguiculata L. Walp) has allowed the investigation of associations between genes of interest and RFLP markers. A cross between an aphid (Aphis craccivora Koch) resistant cultivated cowpea, TT84S-2246-4, and an aphid susceptible wild cowpea, NI 963, was screened for both aphid phenotype and RFLP marker segregation. One RFLP marker, bg4D9b, was found to be tightly linked to the aphid resistance gene (Rac ₁) and several flanking markers in the same linkage group (linkage group 1) were also identified. The close association of Rac ₁ and RFLP bg4D9b presents a real potential for cloning this insect resistance gene.     

Variation in seed proteins from mutagen-treated cultivars and selected lines of Vigna unguiculata L. Walp

The storage protein profiles of the seeds of two IITA cowpea cultivars (‘IT84E–124’ and ‘Vita 7’) exposed to three mutagens—sodium azide (NaN₃), ethyl methane sulphonate (EMS) and ⁶⁰Co gamma rays—and those of 18 selected M₃ lines were investigated by sodium dodecyl sulphate (SDS)-polyacrylamide gel electrophoresis (PAGE). The total seed protein, globulin and albumin fractions showed differences in number and intensity of subunit bands, but the least differences were found in the albumin fraction. Both high and low molecular weight bands were observed, the highest being 94.5 kDa and the lowest 12.0 kDa.‘Vita T showed less variability compared with ‘IT84E–124’, as indicated by the relative similarity indices (S.I.) of the two cultivars. The lowest S.I. of 0.200 was found among ‘IT84E–124’ lines while the lowest S.I. among ‘Vita T lines was 0.4706. A number of lines with particular traits were found to be characterized by the presence of specific polypeptide bands. This study demonstrates that induced mutation could create additional variability to supplement existing germplasm and that SDS-PAGE is a useful tool for discriminating and estimating genetic similarities among selections.     

Genetic analysis of Fusarium wilt resistance in cowpea (Vigna unguiculata Walp.)

This study investigated the inheritance of resistance to Fusarium oxysporum f.sp. tracheiphilum (Fot) in cowpea lines. Resistant and susceptible cowpea lines were crossed to develop F₁, F₂ and backcross populations. Reaction to Fot was evaluated in 2015 and 2016 using seed soak and modified root‐dip inoculation methods. The expression of resistance reaction in the F₁ and segregation in F₂ generations indicated the role of dominant gene controlling Fot in cowpea. These results were further supported by the result of backcross (BC₁P₁F₁ and BC₁P₂F₁) progeny tests. The backcross of F₁ with the resistant parent produced progeny that were uniformly resistant, whereas backcross of F₁ with the susceptible parent produced progeny that segregated into 1:1 ratio. The F₂ segregation ratio in the reciprocal cross showed no evidence of maternal effect in the inheritance of the resistance. Allelism test suggests that the gene for resistance in TVu 134 was the same in TVu 410 and TVu 109‐1. We also identified an SSR marker, C13‐16, that cosegregated with the gene conferring resistance to Fot in cowpea.     

Inheritance of yield components and their correlation with yield in cowpea (Vigna unguiculata (L.) Walp.)

Summary The inheritance of yield components was studied in a cross of two varieties of cowpea (Vigna unguiculata), and the relationships among the components and yield were examined in the F₂ of a cross and in a trial of 22 varieties. The main objective was to determine the suitability of using the components as selection criteria for increasing seed yield. All the components appeared to be under polygenic control and transgressive segregation in the F₂ was observed for pod length and number of seeds per pod. Narrow sense heritability estimates ranged from 19.8% for number of pods per plant to 60.3% for pod length. Most of the genotypic correlations between pairs of yield components were significant. Correlation of yield per plant was negative with pod length, but positive with the other components. Number of pods per plant was consistently correlated with yield. However because of low heritability for number of pods per plant, this component may only be used as a preliminary selection criterion while the final selection is based on yield itself.     

The effectiveness of early generation (F3) yield testing and the single seed descent procedures in two cowpea (Vigna unguiculata (L.) Walp.) crosses

Summary Two breeding procedures were compared in two cowpea crosses. Both procedures were started from the same selected F₂ plants in each cross. For the early generation yield testing, the F₃ lines were yield tested and a pedigree and bulk methods followed in F₄ and F₅. Each F₅ line was bulked to provide seed for a yield test in F₆. In the Single Seed Descent (SSD) procedure, a single seed was taken from each F₂ plant to produce the F₃ generation. The procedure was repeated for the F₄ and F₅. The F₆ SSD lines and the F₆ yield testing lines were compared in yield tests.The results showed that differences in yielding ability of F₃ lines persisted over generations indicating that selection was effective. This was confirmed by the high significant correlations between F₃ yields and those of later generations which ranged from r=0.51^* to 0.85^**.The grain yields of lines derived by the single seed descent procedure were as good as those derived from early generation yield testing.Significant linear correlation between visual rating of F₃ and F₆ yields with actual yields revealed that it is possible to identify promising lines of cowpea visually.     

Isolation, sequence analysis, and linkage mapping of resistance-gene analogs in cowpea (Vigna unguiculata L. Walp.)

Degenerate oligonucleotides designed to recognize conserved coding regions within the nucleotide binding site (NBS) and hydrophobic region of known resistance (R)genes from various plant species were used to target PCR to amplify resistance gene analogs (RGAs) from a cowpea (Vigna unguiculata L. Walp.) cultivar resistant to Striga gesnerioides. PCR products consisted of a group of fragments approximately 500 bp in length that migrated as a single band during agarose gel electrophoresis. The nucleotide sequence of fifty different cloned fragments was determined and their predicted amino acid sequences compared to each other and to the amino acid sequence encoded by known resistance genes, and RGAs from other plant species. Cluster analysis identified five different classes of RGAs in cowpea. Gel blot analysis revealed that each class recognized a different subset of loci in the cowpea genome. Several of the RGAs were associated with restriction fragment length polymorphisms, which allowed them to be placed on the cowpea genomic map. The potential for using these sequences to isolate R genes, and subsequent direct manipulation of disease and pest resistance using genetic engineering is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Evaluation of Pedigree and Single Seed Descent Selection Methods for Cultivar Development in Cowpea (Vigna unguiculata L. Walp)

Three crosses of cowpea (Vigna unguiculata [L.] Walp) were used to assess the effectiveness of two selection procedures in cowpea. The selection procedures were pedigree selection (PD) and single seed descent selection (SSD). Lines developed through each procedure were yield tested and selected at F₆ and F₇ for PD and SSD, respectively. The selected lines for both procedures were compared at F₈ for grain yield per plant (YLD), number of pods per plant (NPO), and pod development period (PDP), in two locations. Lines developed through both procedures differed more in their population means than in the performance of their top 10% or highest yielding lines. There was no consistent relationship between magnitude of genetic variability and percentage of superior lines obtained from each procedure. Both procedures have been effective in producing superior genotypes for yield and number of pods. The PD method produced superior transgressive segregants in two out of the three crosses for PDP. SSD allowed a more rapid generation than PD.     

Effect of Water Stress on the Yield of Cowpea (Vigna unguiculata L. Walp.) Genotypes in the Delmarva Region of the United States

Drought is an important yield-reducing factor for corn and soya bean which are the two major crops in the Delaware, Maryland and Virginia (Delmarva) region of the United States. Cowpea (Vigna unguiculata L. Walp.) is primarily grown in drier regions of the world where it is one of the most drought-resistant food legumes. Field experiments were conducted in which 10 genetically diverse cowpea genotypes were evaluated for adaptability to the Delmarva area. The cowpea genotypes were grown in rain-out shelters under non-water-stressed and water-stressed conditions. The results showed that under non-water-stressed conditions cowpea genotypes California Blackeye 5, Champion and Mississippi Silver gave higher seed yields, while genotypes White Acre, Six Week Browneye and Texas Cream 8 provided lower seed yields. Genotypes California Blackeye 5 and Champion gave comparatively better seed yields under water-stressed conditions. California Blackeye 5 was the highest seed-yielding genotype under both water-stressed and non-water-stressed conditions. The highest biological yield under non-water-stressed conditions was given by genotypes Two Crop Brown, White Acre and Elite, whereas under the water-stressed condition genotypes Texas Cream 8, California Blackeye 5, and Mississippi Silver gave higher biological yield. Genotypes Quickpick Pinkeye and Elite were identified as early maturing genotypes. The harvest index (HI) varied significantly among genotypes, with Texas Cream 8 having the lowest HI. Cowpea genotypes which gave higher seed yield under water-stressed conditions could play an important role in sustaining crop production in the Delmarva region.     

Genetic characterization of Malawian cowpea (Vigna unguiculata (L.) Walp) landraces: diversity and gene flow among accessions

Cowpea has been cultured from ancient timesin Africa and Asia and is now widespreadthroughout the tropics and subtropics.Cowpea accessions can be distinguishedphenotypically from one another by theirgrowth habit, time to maturity, yield, andseed size and colour. Data on geneticdiversity of farmer-developed accessionsare lacking. The main objective of thisstudy was to determine the pattern andextend of RAPD marker variation within andamong cowpea populations from differentagroecological zones and to determine thedegree of genetic relationships and geneflow among different landraces used bylocal farmers. Twenty of the 30 RAPDprimers tested allowed amplifications ofrandom polymorphic (RAPD) loci. Overall,80% of the scored loci were polymorphic. The degree of band sharing was used toevaluate genetic distance betweenaccessions and to construct a phylogenetictree. The genetic distance values amongaccessions varied between 0.09 to 0.57.Although some small clusters groupedaccessions of the same growth habits, ageneral lack of agreement betweenclustering and morphological features wasobserved. The analysis of molecularvariance revealed that within-region ortypes (among accessions) variationaccounted for 96% of the total molecularvariance. This high within-accessionvariability is being sustained by anuncontrolled gene flow among populations.