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In the West African sub-region, intercropping is a critical element in cowpea product profiles. This study aimed to investigate the impact of intercropping cowpea with pearl millet on genetic correlations, heritability, and response to selection. Thirty-five cowpea genotypes and one pearl millet variety were used to create three cropping patterns: sole cowpea, 1:1 (one row of millet to one row of cowpea), and 2:4 (two rows of millet to four rows of cowpea). Split-plot experiments were conducted for two years with cropping patterns considered as the main plots and genotypes as sub-plots. Data were captured on grain yield (GY), 100 seed weight (Hsdwt), pod weight (Pdwt), fodder weight (Fdwt), and harvest index (HI). Genotype, cropping pattern, year, and genotype-by-cropping pattern interactions were significant for most of the traits. Higher heritability and predicted genetic advance were depicted under sole cropping rather than in the intercropped systems. High positive genetic correlations (rg ≥ 0.9) were obtained between cropping patterns. Genetic correlations among traits were higher under sole cropping than in the intercropping. Path analysis portrayed the HI as having the highest significant and positive direct effect on GY. This study identified short duration cowpea genotypes that are stable and adapted to unique cropping systems.

Gender-sensitive cowpea market segments were identified for Nigeria. For the case of cowpea in Nigeria, market segments for producers should also capture consumer preferred traits, given that most of the cowpea consumed within the country is also produced within. A combination of hedonic pricing method and a choice experiment were used to identify gender-sensitive market segments for cowpea in Nigeria. The hedonic pricing method was used to identify key consumer and producer traits (including important hidden traits) to use in the choice experiment. From the choice experiment, 8 market segments were identified across Target Production Environments (TPEs) in Nigeria. These market segments imply that white, brown and speckled cowpea are consumed in Nigeria; also, the largest market segment tends to prefer both large and small/medium cowpea varieties. Cowpea with large size is usually preferred for meals where the cowpea grain is seen (bean porridge, etc.), whereas cowpea with small/medium size is preferred for food involving cowpea paste/flour. The results from this study will support the development of product profiles for Nigeria; they will also help to identify game changers (market segments) for Nigeria., {"references":["Taiwo, Kehinde A. \"The potential of cowpea as human food in Nigeria.\" Food reviews international 14, no. 4 (1998): 351-370.","Gbègbèlègbè, S. D., J. Lowenberg‐DeBoer, R. Adeoti, Jayson Lusk, and O. Coulibaly. \"The estimated ex ante economic impact of Bt cowpea in Niger, Benin and Northern Nigeria.\" Agricultural economics 46, no. 4 (2015): 563-577.","Manda, Julius, Arega D. Alene, Adane H. Tufa, Tahirou Abdoulaye, Tesfamicheal Wossen, David Chikoye, and Victor Manyong. \"The poverty impacts of improved cowpea varieties in Nigeria: A counterfactual analysis.\" World Development 122 (2019): 261-271."]}

Cowpea aphids (

Cowpea (Vigna unguiculata (L.) Walp) is susceptible to several viruses in West Africa. Cowpea viral diseases are mainly controlled through the use of resistant cultivars. Co-infection with more tha...

Management and distribution of experimental data from prebreeding projects is important to ensure uptake of germplasm into breeding and research programs. Being able to access and share this data in standard formats is essential. The adoption of a common informatics platform for crops that may have limited resources brings economies of scale, allowing common informatics components to be used across multiple species. The close integration of such a platform with commonly used breeding software, visualization, and analysis tools reduces the barrier for entry to researchers and provides a common framework to facilitate collaborations and data sharing. This work presents significant updates to the Germinate platform and highlights its value in distributing prebreeding data for 14 crops as part of the project ‘Adapting Agriculture to Climate Change: Collecting, Protecting and Preparing Crop Wild Relatives’ (hereafter Crop Trust Crop Wild Relatives project) led by the Crop Trust (https://www.cwrdiversity.org). The addition of data on these species compliments data already publicly available in Germinate. We present a suite of updated Germinate features using examples from these crop species and their wild relatives. The use of Germinate within the Crop TrustCropWildRelatives project demonstrates the usefulness of the system and the benefits a shared informatics platform provides. These data resources provide a foundation on which breeding and research communities can develop additional online resources for their crops, harness new data as it becomes available, and benefit collectively from future developments of the Germinate platform.

The variable cowpea productivity across different environments demands evaluating the performance of genotypes in a breeding program prior to their release. The aim of this study was to assess yield stability of eight cowpea advanced breeding lines selected from participatory varietal selection in multilocational trials, and to identify mega-environments for cowpea production in Ghana. The genotypes were evaluated across five environments in 2016 and 2017 in randomized complete block design with three replications. The GEA-R version 4.0 software was used for genotype main effect plus genotype by environment interaction (GGE) biplot analyses. Analysis of variance (PROC GLM of SAS using a RANDOM statement with the TEST option) detected significant variations for location, year, genotype, environment, and their interactions. The results showed that the yield performances of the cowpea genotypes were highly influenced by genotype × environment interaction effects. The principal component 1 (PC1) and PC2 were significant components which accounted for 46.75% and 22.84% of GGE sum of squares, respectively. We showed for the first time, two mega-environments for cowpea production and testing in the major cowpea production agro-ecologies in Ghana. The genotypes SARI-6-2-6 and IT07K-303-1 were adapted to Damongo, Nyankpala, and Tumu, whereas SARI-2-50-80 was adapted to Yendi and Manga. The best ranking location was Damongo followed by Tumu, and Nyankpala. The high-yielding genotypes, IT86D-610, IT10K-837-1, IT07K-303-1, and SARI-2-50-80 had significant higher grain yields than the check (Bawutawuta) and were recommended for release as cultivars (or as breeding lines) to boost cowpea production in Ghana.

SummaryComparative analysis of preferences and key criteria for selecting cowpea (Vigna unguiculata L. Walp.) varieties was conducted using the data collected from farmers’ participatory varietal selection (FPVS) activities conducted over 3 years with a total of 2401 farmers (1230 male and 1171 female) in 13 villages in Tougouri department in northern region and Tiéfora department in southern region of Burkina Faso in West Africa. Over the 3 years, farmer criteria for variety selection remained basically stable, but some variations among the regions and years were noticed. Grain yield was the most common and the most important criterion for farmers’ choice in both regions. Farmers in Tougouri (north) put more emphasis on early maturity (90%) and drought resistance (19.7%) as selection criteria than farmers in Tiéfora (south). Farmers in Tiéfora placed statistically significant importance on seed colour and plant type, while farmers in Tougouri did not, and for these selection criteria, there were only slight differences between genders in both areas. Results of stepwise multiple regression indicated that maturity and seed colour in the north, and seed size and seed colour in the south were the most important selection factors for farmers to select cowpea varieties. Improved varieties should have sufficiently good yield to be accepted, but other favoured traits may differ by target region as a reflection of local and regional market demands as well as deep-rooted cultural preferences. Understanding local and regional differences in selection criteria for cowpea varieties is necessary to improve the acceptance of newly released improved varieties. Preferences identified in the participatory activities could inform further development of cowpea breeding strategies for north and south regions of Burkina Faso.

Three hundred and ninety accessions comprising 260 cultivated and 130 wild cowpea accessions were evaluated phenotypically using 27 cowpea descriptors. Morphological evaluation of some qualitative traits revealed 11.92% and 29.23% presence of pigmentation on the stem, 1.53% and 20.76% presence of stripes on the pod, and 0% and 20% presence of hairiness on the plant of cultivated and wild cowpeas respectively. As for the molecular analysis, sixteen SSR primers were employed for genotyping 48 accessions from both wild and cultivated cowpeas. The data generated a dendrogram with three clusters, two of which consisted of wild cowpea while the third cluster comprised all the cultivated cowpeas, including the yard-long-bean (Vigna unguiculata subsp. sesquipedalis) and Vigna unguiculata subsp. cylindrica accessions. Two wild accessions of subsp. dekindtiana, and one each of subsp. kgalagadensis and protracta clustered with cultivated cowpea indicating their relationships with cultivated cowpea, but not with other wild cowpeas. The numbers of polymorphic SSR bands in cultivated and wild cowpeas were 38 and 54, respectively, while the PIC values were 4.47 and 6.14, respectively, showing a greater genetic diversity in wild than in cultivated cowpeas. The subsp. dekindtiana had the highest number (80%) of shared SSR bands with cultivated cowpea followed by subsp. protracta with 54% of shared bands. Five species of wild cowpea have hairs and so could be used in breeding for resistance to insects.

Crop genetic diversity is a sine qua non for continuous progress in the development of improved varieties, hence the need for germplasm collection, conservation and characterization. Over the years, cowpea has contributed immensely to the nutrition and economic life of the people in Togo. However, the bulk of varieties grown by farmers are landraces due to the absence of any serious genetic improvement activity on cowpea in the country. In this study, the genetic diversity and population structure of 255 cowpea accessions collected from five administrative regions and the agricultural research institute of Togo were assessed using 4600 informative diversity array technology (DArT) markers. Among the regions, the polymorphic information content (PIC) ranged from 0.19 to 0.27 with a mean value of 0.25. The expected heterozygosity (He) varied from 0.22 to 0.34 with a mean value of 0.31, while the observed heterozygosity (Ho) varied from 0.03 to 0.07 with an average of 0.05. The average inbreeding coefficient (FIS) varied from 0.78 to 0.89 with a mean value of 0.83, suggesting that most of the accessions are inbred. Cluster analysis and population structure identified four groups with each comprising accessions from the six different sources. Weak to moderate differentiation was observed among the populations with a genetic differentiation index varying from 0.014 to 0.117. Variation was highest (78%) among accessions within populations and lowest between populations (7%). These results revealed a moderate level of diversity among the Togo cowpea germplasm. The findings of this study constitute a foundation for genetic improvement of cowpea in Togo.

Flower bud thrips is one of the most destructive insect pests of cowpea in sub-Saharan Africa. Information on genetic variability among cowpea germplasm and interrelationships among traits under thrips infestation would facilitate the development of resistant varieties. The objectives of the study were to assess genetic variability for thrips resistance, estimate heritability of yield and other traits and investigate inter-trait relationships under thrips infestation. One hundred and fifty-six cowpea lines, including one resistant and one susceptible check, were screened for resistance under natural infestation at two locations in Nigeria, in 2016. Test lines were scored for thrips damage weekly for three consecutive weeks, after removal of spreader plants, to obtain damage scores (DS) 1, 2 and 3 while data were collected on agronomic traits. The data were subjected to analysis of variance from which genetic components of the phenotypic variance were computed. Interrelationships among traits were determined using phenotypic and genotypic correlation, and sequential path analyses. Significant variability was observed among test lines. Lines TVu 6824 and TVNu 1307 were identified as possessing thrips resistance. DS3 had significant genetic and phenotypic correlations with DS1, DS2 and yield-related traits. Number of pods per peduncle, number of peduncles per plant and DS3 were identified as first-order traits. Heritability estimates ranged from 0.55 to 0.73. Genetic variability among the lines suggests the possibility of genetic control of thrips while number of pods per peduncle, number of peduncles per plant and DS3 would serve as useful selection criteria for thrips resistance.

This study determined the gene effects involved in the inheritance of pod length and other yield-related traits and relationships among traits in crosses between two cowpea lines (TVu 2280 and TVu 2027) and a yard-long bean (TVu 6642) line with long pods. Plants of six generations (P1, P2, F1, F2, BC1P1, and BC1P2) derived from TVu 2280 × TVu 6642 and TVu 2027 × TVu 6642 were evaluated under field conditions. Data collected on 14 yield components of each cross were used for generation mean analysis. Gene effects and their magnitudes varied with the crosses, digenic epistatic gene effects were detected for 10 traits in TVu 2280 × TVu 6642 and 11 traits in TVu 2280 × TVu 6642. Only additive gene effect was significant for pod length in TVu 2280 × TVu 6642 while additive, dominant, and two of the three digenic epistatic gene effects were significant in TVu 2027 × TVu 6642. Models that incorporated only significant additive, dominant, and digenic epistasis were adequate for all 14 traits in TVu 2280 × TVu 6642 and eight of the 12 traits in TVu 2027 × TVu 6642 for which model-fitting was possible. Across segregating generation of the two crosses, pod length (PodLNT) was significantly (p &lt, 0.001) correlated with three major yield components viz. pod weight (0.84, 0.77), number of seeds per pod (0.41, 0.30) and seed weight per pod (0.61, 0.29). Significant correlation of PodLNT with seed yield per plant was moderate and significant (p &lt, 0.01–0.001) in the BC1P1 of the two crosses (0.31 and 0.41). An improvement in cowpea seed yield is feasible through selection for long pods in segregating generations involving crosses with yard-long bean.

Incorporation of new sources of genetic diversity into plant breeding programs is crucial for continuing to improve yield and quality, as well as tolerance to abiotic and biotic stresses. A minicore (the “UCR Minicore”) composed of 368 worldwide accessions of cultivated cowpea has been assembled, having been derived from the University of California, Riverside cowpea collection. High-density genotyping with 51,128 SNPs followed by principal component and genetic assignment analyses identified six subpopulations in the UCR Minicore, mainly differentiated by cultivar group and geographic origin. All six subpopulations were present to some extent in West African material, suggesting that West Africa is a center of diversity for cultivated cowpea. Additionally, population structure analyses supported two routes of introduction of cowpea into the U.S.: (1) from Spain to the southwest U.S. through Northern Mexico, and (2) from Africa to the southeast U.S. via the Caribbean. Genome-wide association studies (GWAS) of important agronomic traits including flowering time, resulted in the identification of significant SNPs for all traits and environments. The mapping resolution achieved by high-density genotyping of this diverse minicore collection allowed the identification of strong candidate genes, including orthologs of the ArabidopsisFLOWERING LOCUS T.In summary, this diverse, yet compact cowpea collection constitutes a suitable resource to identify loci controlling complex traits, consequently providing markers to assist with breeding to improve this crop of high relevance to global food and nutritional security.

The objective of this study was to determine genetic potentials in eight sets of cowpea lines for grain yield (GY), hundred seed weight (HSDWT) and days to 50% flowering (DT50FL). A total of 614 F6 genotypes constituting the sets, grouped by maturity, were evaluated across two locations in Northern Nigeria, in an alpha lattice design, two replications each. Data were recorded on GY, HSDWT and DT50FL.Variance components, genotypic coefficient of variation (GCV), and genetic advance (GA) were used to decode the magnitude of genetic variance within and among sets. Genetic usefulness (Up) which depends on mean and variance to score the genetic merits in historically bi-parental populations was applied to groups of breeding lines with mixed parentage. Principal component analysis (PCA) was used to depict contribution of traits to observed variations. GY and DT50FL explained the variance within and between sets respectively. Genotypes were significantly different, although genotype-by-location and set-by-location interaction effects were also prominent. Genetic variance (δ2G) and GCV were high for GY in Prelim2 (δ2G = 45,897; GCV = 19.58%), HSDWT in Prelim11 (δ2G = 7.137; GCV = 17.07%) and DT50F in Prelim5 (δ2G = 4.54; GCV = 4.4%). Heritability varied among sets for GY (H = 0.21 to 0.57), HSDWT (H = 0.76 to 0.93) and DT50FL (H = 0.20 to 0.81). GA and percentage GA (GAPM) were high for GY in Prelim2 (GAPM = 24.59%; GA = 269.05Kg/ha), HSDWT in Prelim11 (GAPM = 28.54%; GA = 4.47 g), and DT50F in Prelim10 (GAPM = 6.49%; GA = 3.01 days). These sets also registered high values of genetic usefulness, suggesting potential application in non-full sib populations. These approaches can be used during preliminary performance tests to reinforce decisions in extracting promising lines and choose among defined groups of lines.

The narrow base of genetic diversity characteristic of cowpea can be attributed to it being self-pollinating, evolving from narrow wild germplasm and exhibiting very limited gene flow between wild and cultivated types. Backcrossing to introduce simply inherited desirable traits and utilization of improved breeding lines and varieties as parents in crossing programs further narrowed the genetic base of cowpea varieties. In most cowpea breeding programs, genes for resistance and market traits were pyramided into lines characterized by high levels of acceptance to farmers and consumers. Besides predisposing widely distributed improved varieties to genetic vulnerability, a narrow base of genetic variation may be contributing to the plateauing in cowpea grain yield, which compromises genetic gains. Cross compatible wild relatives have not been used in variety development because breeders shy away from them due to their tiny seed size, unattractive seed coat color and texture, pod shattering, and susceptibility to viruses. A number of wild cowpea relatives, both within and outside section Catiang of Vigna species, have been evaluated for their reaction to cowpea insect pests and diseases. Vigna vexillata lines were resistant to the legume pod borer (Maruca vitrata), the cowpea weevil (Callosobruchus maculatus), and Striga gesnerioides but are cross incompatible with cultivated cowpea. Some lines among the cross compatible wild relative V. unguiculata ssp. dekindtiana were found to be resistant to aphid in the seedling stage, while others showed good levels of drought and heat tolerance. Molecular markers are being generated to identify quantitative trait loci (QTL) with effects on some desirable attributes in cowpea. Modern breeding tools, including transgenics, can be applied for the improvement of cowpea, bypassing the natural barriers of traditional breeding. Transgenic cowpea with Bt gene cry1Ab showing resistance to M. vitrata has been released in Nigeria. Genome editing, a powerful emerging tool, can also be used for developing improved cowpea varieties with durable resistance to pests and diseases.

Both cowpea and yard-long bean belong to

Both cowpea and yard-long bean belong to Vigna unguiculata ssp. unguiculata but have diverged through human induced evolution in sub-Saharan Africa and Asia, respectively. To map the quantitative trait loci (QTLs) for yield associated traits and derive new lines that may combine the attributes of both types, we developed a F2:3 mapping population derived from a cross between cowpea line TVu2185 and yard-long bean line TVu6642. Using DArT markers, a total of 30 QTLs accounting for 1.8-13.0% phenotypic variation was detected for pod and seed traits. Some novel major QTLs for peduncle number per plant (qPeN2.2), pod length (qPoL3), seed breadth (qSB4), length (qSL7.2) and thickness (qST9) identified on chromosomes 2, 3, 4, 7 and 9, respectively, are particularly interesting and need to be validated. Moreover, we confirmed previously reported QTLs for pod length (qPoL8) and 100-seed weight (qSW8) on chromosome 8 and for seed number per pod (qSN9.2) on chromosome 9 suggesting usefulness for marker-assisted-selection purpose. Notably, some QTLs for these traits were clustered especially on chromosomes 5, 7, 8, 9 and 10 indicating the presence of the same QTL or linked loci in these regions. Moreover, the involvement of epistasis was observed for trait expressions, but compared with the main effect QTLs, the phenotypic effects of epistatic-QTLs detected were much less. The present QTL analysis may provide a useful tool for breeders to formulate efficientbreeding strategy for introgression of the desirable alleles for yield related traits in cowpea using molecular markers.

Cowpea (Vigna unguiculata(L. Walp) is an important grain legume for human and livestock nutrition, especially in sub-Saharan Africa. Aphid,Aphis craccivoraKoch (Homoptera: Aphididae), is one of the most widespread and destructive insect pests of cowpea and host-plant resistance is an effective approach to minimize the pest damage at seedling stage. This study was aimed at identifying resistant sources toA. craccivorawithin the cowpea mini core collection, a set of accessions from the largest world cowpea germplasm collection maintained at the International Institute of Tropical Agriculture (IITA). A total of 375 lines including 373 from IITA mini core collection, one resistant (TVu-801) and one susceptible (TVx-3236) checks were evaluated through artificial infestation in screening cages during the seedling stage. In cages, genotypes were planted in single rows containing four plants. They were arranged in an augmented design in which the two checks were sown in individual cages. Scoring for aphid population and damage levels were carried out on individual plants at 7, 14, and 21 days after planting. Advanced bioassays and biochemical analyses were conducted to investigate the mechanism of resistance toA. craccivora. Overall, three genotypes TVu-6464, TVu-1583, and TVu-15445 showed good levels of resistance comparable to the resistant check TVu-801. The HPLC analyses proved that both low sucrose levels in the plant, as well as high levels of kaempferol and quercetin, aglycones of phenolic compounds, were related with high resistance to aphids. The above genotypes with promising levels of resistance toA. craccivorawill be used in cowpea breeding programs to develop improved resistant lines against this pest.

Multi-parent advanced generation inter-cross (MAGIC) populations are an emerging type of resource for dissecting the genetic structure of traits and improving breeding populations. We developed a MAGIC population for cowpea (Vigna unguiculata L. Walp.) from eight founder parents. These founders were genetically diverse and carried many abiotic and biotic stress resistance, seed quality and agronomic traits relevant to cowpea improvement in the United States and sub-Saharan Africa, where cowpea is vitally important in the human diet and local economies. The eight parents were inter-crossed using structured matings to ensure that the population would have balanced representation from each parent, followed by single-seed descent, resulting in 305 F8 recombinant inbred lines each carrying a mosaic of genome blocks contributed by all founders. This was confirmed by single nucleotide polymorphism genotyping with the Illumina Cowpea Consortium Array. These lines were on average 99.74% homozygous but also diverse in agronomic traits across environments. Quantitative trait loci (QTLs) were identified for several parental traits. Loci with major effects on photoperiod sensitivity and seed size were also verified by biparental genetic mapping. The recombination events were concentrated in telomeric regions. Due to its broad genetic base, this cowpea MAGIC population promises breakthroughs in genetic gain, QTL and gene discovery, enhancement of breeding populations and, for some lines, direct releases as new varieties.

Crop genetic diversity is a sine qua non for continuous progress in the development of improved varieties, hence the need for germplasm collection, conservation and characterization. Over the years, cowpea has contributed immensely to the nutrition and economic life of the people in Togo. However, the bulk of varieties grown by farmers are landraces due to the absence of any serious genetic improvement activity on cowpea in the country. In this study, the genetic diversity and population structure of 255 cowpea accessions collected from five administrative regions and the agricultural research institute of Togo were assessed using 4600 informative diversity array technology (DArT) markers. Among the regions, the polymorphic information content (PIC) ranged from 0.19 to 0.27 with a mean value of 0.25. The expected heterozygosity (He) varied from 0.22 to 0.34 with a mean value of 0.31, while the observed heterozygosity (Ho) varied from 0.03 to 0.07 with an average of 0.05. The average inbreeding coefficient (FIS) varied from 0.78 to 0.89 with a mean value of 0.83, suggesting that most of the accessions are inbred. Cluster analysis and population structure identified four groups with each comprising accessions from the six different sources. Weak to moderate differentiation was observed among the populations with a genetic differentiation index varying from 0.014 to 0.117. Variation was highest (78%) among accessions within populations and lowest between populations (7%). These results revealed a moderate level of diversity among the Togo cowpea germplasm. The findings of this study constitute a foundation for genetic improvement of cowpea in Togo.

The extent of genetic diversity and relatedness of cowpea germplasm from East Africa are poorly understood. A set of 13 microsatellites (SSR) and 151 single nucleotide polymorphisms (SNPs) markers were applied to assess the levels of genetic diversity in a sample of 95 accessions of local cowpea germplasm and inbred lines of Vigna unguiculata. The average genetic diversity (D), as quantified by the expected heterozygosity, was higher for SSR loci (0.52) than for SNPs (0.34). The polymorphic information content was 0.48 for SSR and 0.28 for SNP while the fixation index was 0.095 for SSR and 0.15 for SNPs showing moderate differentiation and high gene flow among cowpea accessions from East African countries. The results of data analysis of both SSR and SNP markers showed similar clustering patterns suggesting a substantial degree of association between origin and genotype. Principal coordinate analysis (PCoA) with SSR and SNP markers showed that accessions were grouped into two and three broad groups across the first two axes, respectively. Our study found that SNP markers were more effective than SSR in determining the genetic relationship among East African local cowpea accessions and IITA inbred lines. Based on this analysis, five local cowpea accessions Tvu-13490, Tvu-6378, Tvu-13448, Tvu-16073, and 2305675 were identified to be tightly clustered sharing several common alleles with the drought tolerant variety Danila when analyzed with SSR and SNP markers. The findings will assist and contribute to future genetic diversity studies aimed at the genetic improvement of local Eastern Africa cowpea accessions for improved overall agronomic performance in general and breeding for drought tolerant in particular.

Cowpea is an important crop for food security, as it is a primary source of protein and cash income for farmers in the dry savanna areas of Sub-Saharan Africa. Improved varieties should contain reasonable amounts of protein in the grains, and to ensure this, timely and cost-effective analytical methods need to be developed. In this study, a procedure for estimating the grain nitrogen (protein) content in single seeds using near infrared spectroscopy was developed. Near infrared spectroscopic analysis of single seeds is a useful tool for breeders and food companies. A total of 200 cowpea germplasm accessions with wide variation in grain nitrogen content (2.44 – 5.12%N) were used for calibration. Two kinds of sample processing were performed for seed preparation (intact seed and husked seed). The models developed were validated using data from 88 and 89 genotypes. The calibration models based on both intact and husked seeds showed a high level of consistency (R2 = 0.88 and 0.88) in cross-validation. The calibration models predicted samples in validation sets with an accuracy of r2 = 0.85, RMSEP = 0.17%N, RPD = 2.55 and r2 = 0.88, RMSEP = 0.15%N, RPD = 2.90, for intact and husked seeds, respectively. With these levels of accuracy, the developed predictive models showed that NIR measurements for single seeds could be useful to determine the uniformity of grain quality, or for rapid screening of the nitrogen content in the seeds of cowpea genotypes.

Perennial habit and floral scent are major traits that distinguish domesticated cowpeas from their wild relatives. However, the genetic basis of these two important traits remains largely unknown in cowpea. Plant longevity, a perenniality-related trait, and floral scent, an outcrossing trait, were investigated using a RIL population derived from a cross between a domesticated and a wild cowpea. QTL analysis revealed three significant loci, one on chromosome 8 associated with plant longevity and two, on chromosomes 1 and 11, for floral scent. Genes within the QTL regions were identified. Genes encoding an F-box protein (Vigun08g215300) and two kinases (Vigun08g217000, Vigun08g217800), and involved in physiological processes including regulation of flowering time and plant longevity, were identified within the perenniality QTL region. A cluster of O-methyltransferase genes (Vigun11g096800, Vigun11g096900, Vigun11g097000, Vigun11g097600, and Vigun11g097800) was identified within the floral scent QTL region. These O-methyltransferase cowpea genes are orthologs of the Arabidopsis N-acetylserotonin O-methyltransferase (ASMT) gene, which is involved in the biosynthesis of melatonin. Melatonin is an indole derivative, which is an essential molecule for plant interactions with pollinators. These findings lay the foundation for further exploration of the genetic mechanisms of perenniality and floral scent in cowpea. Knowledge from this study can help in the development of new extended-growth cycle lines with increased yield or lines with increased outcrossing for population breeding.

Cowpea, Vigna unguiculata (L.) Walp., is an important grain legume grown and consumed not only in the dry savannah areas of Sub-Saharan Africa but also in many other tropical and subtropical regions. It provides income, food and nutrition security to millions of people. Several studies have led to a better understanding of the taxonomy of cowpea and its wild relatives. The species diversity, distribution and evolution of cowpea have been intensively explored. The crop is mainly cultivated in intercropping system where its low plant population does not allow the full expression of the yield potential of the cultivars being grown. Considerable challenges affect the production of this crop despite its comparatively better adaptation to harsh environments. The available genetic resources maintained in the different gene banks are being used for the improvement of cowpea. Germplasm diversity and cultivars characterization were conducted in different studies. Sources of resistance/tolerance to key biotic and abiotic stresses are being identified and introgressed genes involved in new breeding lines are being developed. Improvement strategies were developed to address the major constraints to production while also taking consumer preferences into consideration. Breeding approaches of self-pollinated crops were used in the breeding programs. Application of biotechnology has been suggested to address intractable problems. Considerable effort has been made to genetically transform cowpea. Recent development of genomic resources should support the implementation of molecular breeding to complement conventional breeding and to enhance genetic gain. Key elements needed for successful application of molecular breeding tools include the availability of a high-throughput genotyping platform, high-quality consensus genetic maps, improved phenotyping capability and identification of markers closely linked to target traits. Progress is being recorded in many of these areas which should allow the development of modern breeding programs that will result in effective and efficient development of improved resilient cowpea cultivars.

The importance of cowpea, Vigna unguiculata, in human and animal nutrition and sustainability of soil fertility are recognized globally especially in sub-Saharan Africa (SSA) where the crop is mainly produced in the Savanna and the Sahelian agro ecologies. However, cowpea productivity is adversely affected by both biotic (insect pests, diseases, parasitic weeds, nematodes) and abiotic (drought, heat, low soil fertility) constraints. Appreciable progress has been made in the improvement of cowpea for resistance to some biotic stresses particularly diseases such as bacterial blight, ashy stem blight, marcophomina, parasitic weeds like Striga and Alectra and some insects like aphid, leaf and flower thrips among others. There is need for intensifying research activities with focus on improving cowpea resistance to abiotic stresses. As a crop grown commonly in arid regions, cowpea is subjected to seedling stage, midseason and terminal droughts. In the recent past, the amount of rainfall, during the cropping season in the dry savannah regions of SSA, is getting less. Consequently the cropping season is getting shorter occasioned by late commencement or early cessation of the rain. Farmers in the cowpea producing areas of SSA generally have no access to irrigation hence their crops are grown under rain-fed conditions. With the impending higher frequency of drought in the dry savannah region due to climate change, efforts should be made in developing climate resilient cowpea varieties that farmers will grow. Efforts have been made in enhancing tolerance to drought in some improved cowpea varieties using conventional breeding but progress has been slow in this regard. Drought tolerance is a complex trait and many genes are involved in its inheritance. Pyramiding of these genes in improved varieties would therefore, be desirable. Such varieties with pyramided genes are likely to be stable in performance over the years and across several locations in the savannahs. Recent developments in molecular biology could play significant role in the development of such resilient varieties. In a number of crops, molecular markers associated with resistance loci have been identified and are being used in marker assisted breeding. Marker assisted backcrossing (MABC) is the choice when single traits that are simply inherited are to be moved to varieties with superior performance but lacking in the trait being transferred. Also, marker assisted recurrent selection (MARS) has shown promise in accumulating multiple genes in improved varieties of some crops. Some work has been initiated in cowpea on the use of MARS to pyramid resistance to Striga, yield and drought. Results obtained so far show the potential of this method in pyramiding desirable genes in cowpea. As more resources get committed to cowpea research a solid foundation would be established for the generation of molecular tools that should facilitate their routine application to the improvement of the crop.

The flower bud thrips, Megalurothrips sjostedti Trybom (Thysanoptera: Thripidae), is an economically important pest of cowpea in sub‐Saharan Africa. Varietal resistance is the most preferred, environmentally friendly, cost‐effective and sustainable option for controlling this pest. The objective of this study was to identify sources of resistance to M. sjostedti among mini core accessions from the largest world cowpea germplasm collection maintained at the International Institute of Tropical Agriculture (IITA). The study was conducted during the 2015 and 2016 cropping seasons where 365 accessions were screened under field conditions. Each accession was rated visually for thrips damage score, flower abortion rate, number of pods per plant and number of thrips per flower. The resistance levels observed in genotypes TVu8631, TVu16368, TVu8671 and TVu7325 were similar to that of the resistant check “Sanzisabinli” (called Sanzi) during both seasons. In addition, 56 mini core genotypes showed moderate resistance to thrips damage. High heritability values were associated with thrips damage scores at 65 days after planting (0.60), percentage of effective peduncles (0.59), flower bud abortion rate (0.59), number of pods per plant (0.51) and number of peduncles with pods (0.5). The accessions identified with good levels of resistance to flower bud thrips will be used in cowpea breeding programs to develop improved resistant varieties.

Cowpea (Vigna unguiculata L. Walp) is a warm-season legume with a genetically diverse gene-pool composed of wild and cultivated forms. Cowpea domestication involved considerable phenotypic changes from the wild progenitor, including reduction of pod shattering, increased organ size, and changes in flowering time. Little is known about the genetic basis underlying these changes. In this study, 215 recombinant inbred lines derived from a cross between a cultivated and a wild cowpea accession were used to evaluate nine domestication-related traits (pod shattering, peduncle length, flower color, days to flowering, 100-seed weight, pod length, leaf length, leaf width and seed number per pod). A high-density genetic map containing 17,739 single nucleotide polymorphisms was constructed and used to identify 16 quantitative trait loci (QTL) for these nine traits. Based on annotations of the cowpea reference genome, genes within these regions are reported. Four regions with clusters of QTL were identified, including one on chromosome 8 related to increased organ size. This study provides new knowledge of the genomic regions controlling domestication-related traits in cowpea as well as candidate genes underlying those QTL. This information can help to exploit wild relatives in cowpea breeding programs.

Cowpea (Vigna unguiculataL. Walp) is a warm-season legume with a genetically diverse gene-pool composed of wild and cultivated forms. Cowpea domestication involved considerable phenotypic changes from the wild progenitor, including reduction of pod shattering, increased organ size, and changes in flowering time. Little is known about the genetic basis underlying these changes. In this study, 215 recombinant inbred lines derived from a cross between a cultivated and a wild cowpea accession were used to evaluate nine domestication-related traits (pod shattering, peduncle length, flower color, flowering time, 100-seed weight, pod length, leaf length, leaf width and seed number per pod). A high-density genetic map containing 17,739 single nucleotide polymorphisms was constructed and used to identify 16 quantitative trait loci (QTL) for these nine domestication-related traits. Candidate genes underlying each of those 16 QTL were identified. Four regions with clusters of QTL were identified, including one on chromosome 8 related to increased organ size. This study provides new knowledge of the genomic regions controlling domestication-related traits in cowpea as well as candidate genes underlying those QTL. This information can help to exploit wild relatives in cowpea breeding programs.Key messageThis study identified regions of the cowpea genome that played an important role in cowpea domestication, including a hotspot region for increased organ size

Cowpea (Vigna unguiculata L. Walp.) is an important cash, food and nutritional security grain legume crop in the semi-arid regions of sub-Saharan Africa. However, its productivity is hampered by several biotic stress factors including numerous insect pests that infest and damage the crop at all its development stages in the field as well as during storage. Host plant resistance is an environmental-friendly, cost-effective and sustainable pest management option for minimizing the pests’ incidence and severity. This review article aims at describing the major insect pests in cowpea and highlight key past and recent research findings in cowpea resistance to insect pests. It also provides in-depth knowledge in the host-plant resistance mechanisms in cowpea i.e. biophysical, biochemical and physiological factors that regulate the defense systems in the plant. Furthermore, the paper discusses the need for advanced investigation on the genetic basis of the plant defense systems and its application to the crop breeding program for developing new improved materials. The review would support the cowpea breeding program with the overall expectations of developing insect-resistant lines, reducing the input costs of insecticides while also enhancing cowpea productivity in sub-Saharan Africa.

Flower bud thrips, Megalurothrips sjostedti is the most severe field pest of cowpea that causes massive flower abortion which eventually results to substantial yield reduction in Africa. There is paucity of information on the mode of gene actions controlling inheritance of resistance to flower bud thrips in cowpea in the literature. The objectives of study were to assess the genetic variability for thrips resistance among the cowpea germplasm, determined the mode of inheritance of genes that conferred resistance and both broad and narrow-sense heritability estimates for the inheritance of thrips resistance in cowpea. Twelve cowpea lines were used in crosses in the screen house at IITA, Ibadan. The mating was accomplished using North Carolina design II to generate 48F1 hybrids, which were eventually evaluated with the parents. Data on number of peduncles, number of pods and number of thrips per flower were recorded and subjected to analysis of variance using random model by SAS 9.2. Significant variability was observed for most agronomic and thrip-adaptive traits among the cowpea germplasm, parental-lines and F1 genotypes evaluated. General combining ability (GCA) and specific combining ability (SCA) mean squares were significant (P

Development and analysis of Multiparent Advanced Generation Inter-Cross (MAGIC) populations have been conducted with several crop plants to harness the potential for dissecting the genetic structure of traits and improving breeding populations. We developed a first MAGIC population for cowpea (Vigna unguiculataL. Walp.) from eight founder parents which are genetically diverse and carry many abiotic and biotic stress resistance, seed quality and agronomic traits relevant to cowpea improvement in sub-Saharan Africa (SSA) where cowpea is vitally important in the human diet and in local economies. The eight parents were inter-crossed using structured matings to ensure the population would have balanced representation from each of the founder parents, followed by single-seed descent, resulting in 365 F8 recombinant inbred lines (RILs) each carrying a mosaic of genome blocks contributed from all founders. This was confirmed by SNP genotyping with the cowpea Illumina 60K iSelect BeadArray. Following filtering to eliminate duplicates, sister lines and accidental selfing events, a core set of 305 F8 RILs was chosen as the primary population. The F8 lines were on average 99.74% homozygous while also diverse in agronomic traits including flowering time, growth habit, maturity, yield potential and seed characteristics across environments. Trait-associated SNPs were identified for most of the parental traits. Loci with major effects on photoperiod sensitivity and seed size were also verified by genetic mapping in biparental RIL populations. The distribution of recombination frequency varied considerably between chromosomes, with recombination hotspots distributed mostly in the telomeric regions. Due to its broad genetic base, this cowpea MAGIC population promises breakthroughs in genetic gain and high-resolution genetic mapping for gene discovery, enhancement of breeding populations and, for some lines, direct releases as new varieties.

Summary Cowpea (Vigna unguiculata L. Walp.) is a legume crop that is resilient to hot and drought-prone climates, and a primary source of protein in sub-Saharan Africa and other parts of the developing world. However, genome resources for cowpea have lagged behind most other major crops. Here we describe foundational genome resources and their application to analysis of germplasm currently in use in West African breeding programs. Resources developed from the African cultivar IT97K-499-35 include a whole-genome shotgun (WGS) assembly, a bacterial artificial chromosome (BAC) physical map, and assembled sequences from 4,355 BACs. These resources and WGS sequences of an additional 36 diverse cowpea accessions supported the development of a genotyping assay for 51,128 SNPs, which was then applied to five biparental RIL populations to produce a consensus genetic map containing 37,372 SNPs. This genetic map enabled the anchoring of 100 Mb of WGS and 420 Mb of BAC sequences, an exploration of genetic diversity along each linkage group, and clarification of macrosynteny between cowpea and common bean. The SNP assay enabled a diversity analysis of materials from West African breeding programs. Two major subpopulations exist within those materials, one of which has significant parentage from South and East Africa and more diversity. There are genomic regions of high differentiation between subpopulations, one of which coincides with a cluster of nodulin genes. The new resources and knowledge help to define goals and accelerate the breeding of improved varieties to address food security issues related to limited-input small-holder farming and climate stress. This article is protected by copyright. All rights reserved.

Cowpea is an important legume crop in Africa, valued highly for its grain and also haulms, which are a tradable commodity in fodder markets. Fodder market surveys in Northern Nigeria showed that groundnut haulms were priced higher than cowpea haulms, probably because of their superior nutritive value. The economic value of haulms has prompted cowpea breeders and livestock nutritionists to explore haulm fodder traits as additional selection and breeding criteria. Fifty cowpea genotypes cultivated across five locations in Nigeria in 2013 and 2014 were evaluated for food fodder traits. Significant (P < 0.05) genotypic dependent variations were observed in yields (kg/ha) of grains (537–1082) and haulms (1173–3368), though significant (P < 0.05) effects of location and year were observed. Grain and fodder yield had a tendency to be positively correlated (r = 0.26, P = 0.07). Haulms were analyzed for nitrogen (N), fiber fractions, in vitro digestibility, and metabolizable energy content. Highly significant variations were observed in all genotypic and livestock nutrition traits, although location and year had significant effects. Trade-offs between grain yield and haulm fodder quality traits were largely absent and haulm acid detergent lignin and grain yield were even inversely correlated (r = -0.28, P = 0.05), that is high grain yielders had decreased haulm lignin. However, haulm N and grain yield also tended to be negatively associated (r = -0.26, P = 0.07). Haulm fodder quality traits and haulm yield were mostly positively correlated (P < 0.05). Broad sense heritabilities for grain and fodder yield were 0.50 and 0.29, respectively, while heritability for haulm fodder quality traits ranged from 0.61 to 0.67, providing opportunities for concomitant increase in grain yield and haulm fodder quality traits. Selection of the 10 highest ranking genotypes for grain yield, haulm yield, haulm N, and haulm in vitro organic matter digestibility showed selection groups overlapping, suggesting that multi-trait selection is feasible. Economical evaluation showed that choice of primary traits is context specific, highlighting the need for identifying and targeting appropriate genotypes to fit different production systems. Considering haulm quantity and quality as traits of economic value can increase overall plant value in mixed crop-livestock systems.

SUMMARYCowpea (Vigna unguiculata L. Walp.) is a legume crop that is resilient to hot and drought-prone climates, and a primary source of protein in sub-Saharan Africa and other parts of the developing world. However, genome resources for cowpea have lagged behind most other major crop plants. Here we describe foundational genome resources and their application to analysis of germplasm currently in use in West African breeding programs. Resources developed from the African cultivar IT97K-499-35 include bacterial artificial chromosome (BAC) libraries and a BAC-based physical map, assembled sequences from 4,355 BACs, as well as a whole-genome shotgun (WGS) assembly. These resources and WGS sequences of an additional 36 diverse cowpea accessions supported the development of a genotyping assay for over 50,000 SNPs, which was then applied to five biparental RIL populations to produce a consensus genetic map containing 37,372 SNPs. This genetic map enabled the anchoring of 100 Mb of WGS and 420 Mb of BAC sequences, an exploration of genetic diversity along each linkage group, and clarification of macrosynteny between cowpea and common bean. The genomes of West African breeding lines and landraces have regions of marked depletion of diversity, some of which coincide with QTL that may be the result of artificial selection or environmental adaptation. The new publicly available resources and knowledge help to define goals and accelerate the breeding of improved varieties to address food security issues related to limited-input small-holder farming and climate stress.

Cowpea is one of the most important grain legumes in sub-Saharan Africa (SSA). It provides strong support to the livelihood of small-scale farmers through its contributions to their nutritional security, income generation and soil fertility enhancement. Worldwide about 6.5 million metric tons of cowpea are produced annually on about 14.5 million hectares. The low productivity of cowpea is attributable to numerous abiotic and biotic constraints. The abiotic stress factors comprise drought, low soil fertility, and heat while biotic constraints include insects, diseases, parasitic weeds, and nematodes. Cowpea farmers also have limited access to quality seeds of improved varieties for planting. Some progress has been made through conventional breeding at international and national research institutions in the last three decades. Cowpea improvement could also benefit from modern breeding methods based on molecular genetic tools. A number of advances in cowpea genetic linkage maps, and quantitative trait loci associated with some desirable traits such as resistance to Striga, Macrophomina, Fusarium wilt, bacterial blight, root-knot nematodes, aphids, and foliar thrips have been reported. An improved consensus genetic linkage map has been developed and used to identify QTLs of additional traits. In order to take advantage of these developments single nucleotide polymorphism (SNP) genotyping is being streamlined to establish an efficient workflow supported by genotyping support service (GSS)-client interactions. About 1100 SNPs mapped on the cowpea genome were converted by LGC Genomics to KASP assays. Several cowpea breeding programs have been exploiting these resources to implement molecular breeding, especially for MARS and MABC, to accelerate cowpea variety improvement. The combination of conventional breeding and molecular breeding strategies, with workflow managed through the CGIAR breeding management system (BMS), promises an increase in the number of improved varieties available to farmers, thereby boosting cowpea production and productivity in SSA.