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.     

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.     

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.     

Evaluation of genetic diversity among commercial cultivars, hybrids and local mango (Mangifera indica L.) genotypes of India using cumulative RAPD and ISSR markers

An assessment of genetic diversity studies was undertaken to understand the level and pattern of diversity in 65 mango (Mangifera indica L.) genotypes of India including 20 commercial cultivars, 18 hybrids, 25 local genotypes and two exotic cultivars based on qualitative and quantitative fruit characters as well as RAPD and ISSR profiles. A considerable variation was observed in respect of three important qualitative characters namely table quality, fruit attractiveness and storage life of ripe fruits and potentially superior genotypes for the above traits were identified. A wide variation was noticeable regarding metabolite composition of pulp of ripe mango fruit with respect to total soluble solids, total sugar, reducing sugar, acidity, sugar:acid ratio, ascorbic acid and phenolic content. Fifteen RAPD primers yielded 27 monomorphic and 129 polymorphic bands with percent polymorphism averaging 82.7%. Of a total 70 ISSR bands generated from eight ISSR primers, 60 bands (85.71%) were found to be polymorphic. Cumulative band data from these two methods precisely arranged accessions into eight clusters which correspond well with their pedigree relationship. UPGMA dendrograms drawn using RAPD, ISSR and cumulative data showed highly similar grouping of genotypes on the basis of their parental origin. No clear-cut geographical separation was revealed among East, West, North and South Indian mango cultivars by neither of these molecular markers nor their combinations. This supports the common gene pool origin of mango as well as operation of similar selection pressure as the cultivar preferences in these areas are largely similar.     

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.     

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.     

Molecular characterization and genetic relatedness among walnut (Juglans regia L.) genotypes based on RAPD markers

The potential use of the Randomly Amplified Polymorphic DNA (RAPD) technique for characterization and assessment of genetic relationships was investigated in nineteen walnut (Juglans regia L.) genotypes used as parents or released as cultivars from the breeding program of the University of California at Davis. Most of the 72 decamer primers used yielded scorable amplification patterns based on discernable bands. The results obtained produced a unique fingerprint for each of the walnut genotypes studied. Cluster analysis separated the 19 walnut genotypes into two main groups whose differences were related to their pedigree. Genotypes sharing common parents tend to group together and with at least one of the parents. Thus, RAPD markers can detect enough polymorphism to differentiate among walnut genotypes, even among closely related genotypes, and the genetic similarity based on RAPDs appears to reflect the known pedigree information. RAPD technology can be useful in current walnut breeding programs, allowing the identification of new cultivars as well as the assessment of the genetic similarity among genotypes which will help in selecting the best parents to obtain new genetic combinations. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Identification of markers associated with bacterial blight resistance loci in cowpea [Vigna unguiculata (L.) Walp.]

Cowpea bacterial blight (CoBB), caused by Xanthomonas axonopodis pv. vignicola (Xav), is a worldwide major disease of cowpea [Vigna unguiculata (L.) Walp.]. Among different strategies to control the disease including cultural practices, intercropping, application of chemicals, and sowing pathogen-free seeds, planting of cowpea genotypes with resistance to the pathogen would be the most attractive option to the resource poor cowpea farmers in sub-Saharan Africa. Breeding resistance cultivars would be facilitated by marker-assisted selection (MAS). In order to identify loci with effects on resistance to this pathogen and map QTLs controlling resistance to CoBB, eleven cowpea genotypes were screened for resistance to bacterial blight using 2 virulent Xav18 and Xav19 strains isolated from Kano (Nigeria). Two cowpea genotypes Danila and Tvu7778 were identified to contrast in their responses to foliar disease expression following leaf infection with pathogen. A set of recombinant inbred lines (RILs) comprising 113 individuals derived from Danila (resistant parent) and Tvu7778 (susceptible parent) were infected with CoBB using leaf inoculation method. The experiments were conducted under greenhouse conditions (2007 and 2008) and disease severity was visually assessed using a scale where 0 = no disease and 4 = maximum susceptibility with leaf drop. A single nucleotide polymorphism (SNP) genetic map with 282 SNP markers constructed from the same RIL population was used to perform QTL analysis. Using Kruskall-Wallis and Multiple-QTL model of MapQTL 5, three QTLs, CoBB-1, CoBB-2 and CoBB-3 were identified on linkage group LG3, LG5 and LG9 respectively showing that potential resistance candidate genes cosegregated with CoBB resistance phenotypes. Two of the QTLs CoBB-1, CoBB-2 were consistently confirmed in the two experiments accounting for up to 22.1 and to 17.4% respectively for the first and second experiments. Whereas CoBB-3 was only discovered for the first experiment (2007) with less phenotypic variation explained of about 10%. Our results represent a resource for molecular marker development that can be used for marker assisted selection of bacterial blight resistance in cowpea.     

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.     

Mapping of quantitative trait loci for floral scent compounds in cowpea (Vigna unguiculata L.)

Floral scent is a very important trait in plant evolution. Currently, little is known about the inheritance of floral scent in cowpea (Vigna unguiculata L.) or changes that might have occurred during its domestication. Therefore, we analysed scent volatiles and molecular markers in a population of 159 F₇ recombinant inbred lines derived from a cross of a domesticated blackeye cowpea cultivar, ‘524B’ and a wild accession ‘219‐01’. Using gas chromatography‐mass spectrometry (GC–MS) 23 volatile compounds were identified that fall into five general functional categories. Twenty‐two of the compounds displayed quantitative variation in the progeny, and a total of 63 QTLs influencing the amounts of these volatiles were mapped onto the cowpea genetic marker map. Although QTLs for volatile compounds putatively involved in cowpea flower scent were found on 9 of the 11 cowpea chromosomes, they were not evenly distributed with QTLs mainly clustered on LGs 1, LGs 2 and LG 4. Our results serve as a starting point for both more detailed analyses of complex scent biosynthetic pathways and the development of markers for marker‐assisted breeding of scented rose varieties.     

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.     

Analysis of the genetic diversity among mandarins (Citrus spp.) using RAPD markers

RAPD markers were used to evaluate genetic similarity among 35 mandarin accessions, including 10 species and 7 hybrids. One octamer and twenty-two decamer primers produced 109 RAPDs, 45 of which were polymorphic. Jaccard coefficient was used to calculate genetic similarity, and UPGMA to generate the phenogram. The RAPDs obtained were sufficient to generate some accession-specific markers, and to separate these accessions by clustering them into several groups, many of them according to Tanaka's or Webber's systematic units. The genetic similarity within the mandarin group is high (G_J = 0.77), and suggests that cultivated mandarins have a narrow genetic base. The genetic similarity of mandarins to other true citrus species (Citron [C. medica L.] and Pummelo [C. grandis Osbeck]) was much lower (minimum G_J = 0.27). We propose that the mandarin group is a single species, C. reticulata Blanco, composed of several genetically different individuals and a great number of hybrids, rather than a large number of species as proposed by some taxonomic studies. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

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.     

Identification and genetic diversity analysis of date palm (Phoenix dactylifera L.) varieties from Morocco using RAPD markers

Genetic variation among 43 date palm (Phoenix dactylifera L.) accessions, including 37 accessions from Morocco and 6 cultivars from Iraq and Tunisia, was studied using Random Amplified Polymorphic DNA (RAPD) markers. The pre-screening of 123 primers on four genotypes allowed selection of 19 primers which revealed polymorphism and gave reproducible results. All 43 analysed genotypes were distinguishable by their band patterns. RAPD technology therefore appears very effective for identifying accessions of date palm. RAPD-based genetic distance was used to determine the relationships between the accessions. The grouping-association identified by cluster analysis was rather weak. However, morphologically similar varieties clustered together. A relatively low polymorphism and a lack of evident organisation are observed among the date palm varieties grown in Morocco. This could be related to the mode of introduction and maintenance of the Moroccan date palm germplasm involving limited foundation germplasm, exchange of cultivars between plantations, and periodic development of new recombinant cultivars following sexual reproduction. This revised version was published online in August 2006 with corrections to the Cover Date.     

AFLP, ISSR and RAPD markers reveal high levels of genetic diversity among Lupinus spp.

The Lupinus genus includes a number of important crop species. The use of defined nucleotide sequences for the analysis of genetic diversity among these species has revealed modest levels of diversity. The aim of this study was to access AFLP, ISSR and RAPD markers to evaluate the genetic diversity among L. albus, L. angustifolius, L. cosentinii, L. hispanicus, L. luteus, L. mutabilis, L. pilosus and L. polyphyllus. Unexpectedly, low levels of genetic similarity were found (ranging from 0.205 to 0.432), regardless of the type of molecular marker used. Nevertheless, these techniques consistently showed a greater genetic similarity between L. pilosus and L. cosentinii, L. mutabilis and L. polyphyllus and among L. luteus, L. hispanicus and L. angustifolius, clearly separating the Old World from the New World species. Such low genetic similarity among Lupinus spp. is most unlikely to be due to differences in coding sequences but could be the result of a long diverging process concerning non‐coding regions, which would represent a very important proportion of these genomes.     

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.