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.     

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.     

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.     

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.     

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.     

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.     

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.     

Digestive enzyme inhibitors and storage pest resistance in cowpea (Vigna unguiculata) seeds

Summary Amylase and trypsin inhibitors are proteins which inhibit digestive enzymes. The loss of the activity of these enzymes produces a reduction of starch and protein digestion. Amylase and trypsin inhibitor activity have been investigated in seeds of 20 cowpea lines in trying to establish their role in the storage pest resistance. A broad variation has been observed for both the inhibitors. Correlation analysis between inhibitor levels and extent of attack has shown that neither amylase nor trypsin inhibitors can separately explain the resistance to Callosobruchus maculatus (F.). Lines bruchid resistant in fact, have high levels of both inhibitors. Conversely lines with low levels of amylase and trypsin inhibitors are bruchid susceptible. The breeding for high contents of both amylase and trypsin inhibitors can be an effective way to obtain lines with resistance to storage pest.     

Inheritance of flower and pod colour in cowpea (Vigna unguiculata L. Walp.)

Inheritance of flower colour and pod colour in cowpea (Vigna unguiculata L. Walp.) has followed a qualitative pattern. Purple flower colour is dominant over white flower colour, whereas black pod colour is partially dominant over white pod colour. A segregation ratio of 3 purple:1 white flowers in F₂ generations of two crosses indicated that white flower colour is controlled by a single recessive. Segregation ratio of F₂ 1 white:2 light black:1 black indicated that black pod colour is partially dominant over white pod colour and is governed by one gene. These results were further confirmed by backcross generations. White flower and pod colour are controlled by single recessive genes on separate chromosome. Gene symbols were assigned. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

New sources of resistance of cowpea (Vigna unguiculata) toStriga gesnerioides,a parasitic angiosperm

Summary Thirty-seven accessions of cowpea and yard-long bean were assessed for resistance toStriga gesnerioides. Cowpea plants were grown using anin vitro method, then inoculated with young seedlings ofS. gesnerioides produced from seed from three West African countries. Resistance was assessed by comparing the number and size ofS. gesnerioides tubercles on these accessions with those on a known susceptible cowpea, cv. Blackeye. Two cowpea landraces, APL-1 and 87-2, were completely resistant toS. gesnerioides from Burkina Faso, Mali and Cameroon and partially resistant toS. gesnerioides from Niger. Complete resistance was expressed either as a hypersensitive response of infected root tissues or as a severely retarded development of successful infections. All other accessions, including three samples of yard-long bean were susceptible toS. gesnerioides. The original 87-2 plants segregated for resistance and susceptibility. However, uniformly resistant progeny were obtained by producing seed from vegetatively propagated clones of single resistant 87-2 plants. Resistance of APL-1 and 87-2 toS. gesnerioides was confirmed in pot and field trials. Neither of these cowpeas were resistant toAlectra vogelii. Varieties APL-1 and 87-2 provide additional sources of resistance to most races ofS. gesnerioides, including a newly discovered virulent race from Benin.Abbreviations ICRISAT International Crops Research Institute for the Semi-Arid Tropics - IITA International Institute of Tropical Agriculture - SAFGRAD Semi-Arid Food Grain Research and Development     

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.     

A stability analysis of time to flowering as a screen for responsiveness to temperature and photoperiod in cowpea (Vigna unguiculata)

Summary Twenty-one genotypes of cowpea (Vigna unguiculata), comprising landraces and varieties, were grown in 22 photothermal environments in Nigeria and Niger, West Africa, and a stability analysis of days from sowing to flowering (f) was carried out. Cowpeas are rarely insensitive to photoperiod; they are typically quantitative shortday plants wherein f is delayed when photoperiod (P) is longer than the critical photoperiod (P _c ). Therefore, in order to quantify genotypic variation in temperature sensitivity, genotype f was regressed against the mean trial f in circumstances where P c (i.e. approximately 13 hd^-1) and mean temperature (T) was between 19° and 28° C. Correspondingly, in order to assess genotypic variation in photoperiod sensitivity, trials where T was near optimal (25°–28° C) but where P ranged from 10–14.5 hd^-1 were used. These stability analyses detected no significant differences (P>0.05) between genotypes 9n temperature sensitivity but revealed significant differences (P<0.001) in photoperiod sensitivity. Regression coefficients from the stability analysis were strongly correlated (r=0.94, 19df) with a photoperiod sensitivity constant, c, determined from a photothermal flowering model. A stability analysis of f from field trials can therefore identify and quantify genotypic variation in response to temperature and photoperiod in cowpea.Abbreviations f days from sowing to flowering - P mean photoperiod - P _c critical photoperiod - P _ce ceiling photoperiod - T mean temperature - T _b base temperature - T _o optimum temperature - SDP short-day plant     

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 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.     

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.     

Virulence characteristics of a new race of the parasitic angiosperm,Striga gesnerioides,from southern Benin on cowpea (Vigna unguiculata)

An in vitro growth system was used to determine the virulence of two samples of Striga gesnerioides from Zakpota in southern Benin. Cowpea variety B301, previously considered resistant to all races of S. gesnerioides, was susceptible to both samples of the parasite. Two other cowpea varieties, 58–57 and IT81D-994, were totally resistant. Resistance in 58–57 was associated with a hypersensitive necrosis of infected roots, whilst IT81D-994 supported production of small S. gesnerioides tubercles with stems which failed to develop. Striga gesnerioides from southern Benin is the fourth race of the parasite to be identified, and the first with virulence on variety B301. The implications for breeding cowpeas with resistance to S. gesnerioides are discussed.Abbreviations IITA International Institute of Tropical Agriculture - LARS Long Ashton Research Station - SAFGRAD Semi-Arid Food Grain Research and Development     

Interspecific hybridization between urdbean (Vigna mungo (L.) Hepper) and adzukibean (Vigna angularis (Willd.) Ohwi and Ohashi)

Summary To introduce resistance to fungal and viral diseases into urdbean (Vigna mungo), it was crossed with adzukibean (Vigna angularis) to achieve an interspecific hybrid between the two species by using different non-conventional breeding methods. Partial success was obtained with the use of UV irradiated pollen, intraspecific hybrids as parents and spray of immuno-suppressants on the female parent under glass house conditions and to the extent of callusing in aseptic culture.     

Genetic diversity of wild coffee (Coffea arabica L.) using molecular markers

Genetic diversity was studied using RAPD markers among119 coffee (Coffea arabica L.) individuals representing 88 accessions derived from spontaneous and subspontaneous trees in Ethiopia, the primary centre of species diversity, six cultivars grown locally in Ethiopia, and two accessions derived from the genetic populations Typica and Bourbon, spread in the 18^th century, which gave rise to the most currently grown cultivars. Twenty-nine polymorphic fragments were used to calculate a similarity index and construct dendrograms. The Ethiopian material was separated from the Typica- and Bourbon-derived accessions and classified in four groups: one with most of the collected material from southwestern Ethiopia and three from southern and southeastern Ethiopia. Almost all detected diversity was found in the southwestern group while the southern and southeastern groups presented only 59% of identified markers. The genetic distances were low between the southwestern group and the southern and southeastern groups, and between the southwestern group and the Typica- and Bourbon-derived accessions. The cultivated coffee derived from the genetic populations Typica and Bourbon appeared little differentiated from wild coffee growing in the southwest. The results supported the hypothesis that southwestern Ethiopian coffee trees could have been introduced recently in the south and southeast. A separate analysis of the 80accessions classified in the southwestern group allowed identifying particular spontaneous- and subspontaneous-derived accessions and redundancies in the collected material from southwestern Ethiopia. RAPD markers did not detect any within-collection polymorphism except for two trees that were identified as off-types in the CATIE field genebank. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.