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The role of polyploidy, particularly allopolyploidy, in plant diversification is a subject of debate. Whole-genome duplications precede the origins of many major clades (e.g., angiosperms, Brassicaceae, Poaceae), suggesting that polyploidy drives diversification. However, theoretical arguments and empirical studies suggest that polyploid lineages may actually have lower speciation rates and higher extinction rates than diploid lineages. We focus here on the grass tribe Andropogoneae, an economically and ecologically important group of C4 species with a high frequency of polyploids. A phylogeny was constructed for ca. 10% of the species of the clade, based on sequences of four concatenated low-copy nuclear loci. Genetic allopolyploidy was documented using the characteristic pattern of double-labeled gene trees. At least 32% of the species sampled are the result of genetic allopolyploidy and result from 28 distinct tetraploidy events plus an additional six hexaploidy events. This number is a minimum, and the actual frequency could be considerably higher. The parental genomes of most Andropogoneae polyploids diverged in the Late Miocene coincident with the expansion of the major C4 grasslands that dominate the earth today. The well-documented whole-genome duplication in Zea mays ssp. mays occurred after the divergence of Zea and Sorghum. We find no evidence that polyploidization is followed by an increase in net diversification rate; nonetheless, allopolyploidy itself is a major mode of speciation.

Understanding the genetic basis underly- ing domestication-related traits (DRTs) of cowpea (Vigna unguiculata (L.) Walp.) is important since the genome has experienced divergent domestication and in addition it is also useful to utilize the wild germplasm efficiently for improving different traits of the cultivated cowpea. Quantitative trait loci (QTLs) for DRTs were identified in a population of 159 F7 recombinant inbred lines derived from a cross between a domesticated cowpea (V. unguiculata (L.) Walp.) variety, 524B, and a wild accession, 219. Using the constructed linkage map, QTLs for 10 DRTs were analysed and mapped. QTLs for seed, pod and flower related traits were detected. Subsequently, QTL for ovule number was also identified. To our knowledge, this is the first time a QTL for this trait has been observed. QTLs for DRTs show co-localization on three linkage groups and pleiotropy or close linkage of genes for the traits is suggested in these chromosome regions. The information gained in this study can be used for marker-assisted selection of domestication- related QTLs in cowpea and enhance understanding of domestication in the genus Vigna.

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. (Less)

Flowering time is a major adaptive trait in plants and an important selection criterion in the breeding for genetic improvement of crop species. QTLs for the time of flower opening and days to flower were identified in a cross between a short duration domesticated cowpea (Vigna unguiculata (L.) Walp.) variety, 524B, and a relatively long duration wild accession, 219-01. A set of 159 F7lines was grown under greenhouse conditions and scored for the flowering time associated phenotypes of time of flower opening and days to flower. Using a LOD threshold of 2.0, putative QTLs were identified and placed on a linkage map consisting of 202 SSR markers and four morphological loci. A total of five QTLs related to the time of flower opening were identified, accounting for 8.8%–29.8% of the phenotypic variation. Three QTLs for days to flower were detected, accounting for 5.7%–18.5% of the phenotypic variation. The major QTL of days to flower and time of flower opening were both mapped on linkage group 1. The QTLs identified in this study provide a strong foundation for further validation and fine mapping for developing an efficient way to restrain the gene flow between the cultivated and wild plants.

In 1992, Vaillancourt and Weeden discovered a very important mutation for studying cowpea evolution and domestication. A loss of a BamHI restriction site in chloroplast DNA characterized all domesticated accessions and a few wild (Vigna unguiculata ssp. unguiculata var. spontanea) accessions. In order to screen a larger number of accessions, primers were designed to check this mutation using PCR RFLP or direct PCR methods. Using these new primers, 54 domesticated cowpea accessions and 130 accessions from the wild progenitor were screened. The absence of haplotype 0 was confirmed within domesticated accessions, including primitive landraces from cultivar-groups Biflora and Textilis, suggesting that this mutation occurred prior to domestication. However, 40 var. spontanea accessions distributed from Senegal to Tanzania and South Africa showed haplotype 1. Whereas this marker could not be used to identify a precise center of origin, it did highlight the widely distributed cowpea crop-weed complex. Its very high frequency in West Africa could be interpreted as a result of either genetic swamping of the wild/weedy gene pool by the domesticated cowpea gene pool or as the result of domestication by ethnic groups focusing primarily on cowpea as fodder.

Evidence from chloroplast trnK and nuclear ribosomal ITS sequences and morphological data reveals that the monotypic legume genus Wajira is nested within a clade comprising the species of Vigna subgen. Macrorhynchus. This Wajira-containing clade is basally branching in a larger clade that contains many of the genera traditionally referred to as tribe Phaseoleae subtribe Phaseolinae. Wajira is thus recircumscribed to include Vigna subgen. Macrorhynchus. Given the heterogeneity of floral morphology of its constituent species, Wajira is apomorphically diagnosed by woody stems and a pollen brush that comprises an introrse linear array of unicellular hairs. This recircumscribed genus now comprises five species, one of which is described as new, Wajira danissana. Three species require new nomenclatural combinations, Wajira grahamiana, Wajira praecox, and Wajira virescens. Wajira albescens, W. danissana, W. praecox, and W. virescens are woody climbers that are each narrowly distributed in the arid Somal...

The present study, using RAPD analysis, was undertaken to characterize genetic variation in domesticated cowpea and its wild progenitor, as well as their relationships. The materials used consisted of 26 domesticated accessions, including accessions from each of the five cultivar-group, and 30 wild/weedy accessions, including accessions from West, East and southern Africa. A total of 28 primers generated 202 RAPD bands. One hundred and eight bands were polymorphic among the domesticated compared to 181 among wild/weedy cowpea accessions. Wild accessions were more diverse in East Africa, which is the likely area of origin of V. unguiculata var. spontanea. Var. spontanea is supposed to have spread westward and southward, with a loss of variability, loss counterbalanceed in southern Africa by introgressions with local perennial subspecies. Although the variabilty of domesticated cowpea was the highest ever recorded, cultivar-groups were poorly resolved, and several results obtained with isozyme data were not confirmed here. However primitive cultivars were more diverse than evolved cultivars, which still suggests two consecutive bottlenecks within domesticated cowpea evolution. As isozymes and AFLP markers, although with a larger number of markers, RAPD data confirmed the single domestication hypothesis, the gap between wild and domesticated cowpea, and the widespread introgression phenomena between wild and domesticated cowpea.

A survey of allozyme variation in Kersting's groundnut (Macrotyloma geocarpum(Harms) Marechal and Baudet) was undertaken by examining 19 enzymes systems encoding 32 putative loci in 18 domesticated accessions and two wild accessions. No variation was found within and among domesticated accessions as well as within and between both wild accessions. How- ever, very high genetic distance between wild and domesticated accessions suggests that they should be assigned to two different species. 2002 Elsevier Science Ltd. All rights reserved.

A survey of allozyme variation in cultivar-groups of cowpea [Vigna unguiculata (L.) Walp.] was undertaken by examining 21 enzyme systems encoded by 36 loci in 271 accessions representing the five cultivar-groups. Very low levels of variation were found within accessions, which is typical of self-pollinating species. Little variation was also found among accessions. Compared with other legume crops, V. unguiculata is depauperate in allozyme variation. We found an average of 1.61 alleles per locus with 42% of the loci polymorphic and a total heterozygosity of 0.061. Of the variation present, 90% was found within cultivar-groups, while 10% was among cultivar-groups. Data analyses revealed continuous variation among cultivar-groups and geographic regions with the accessions failing to segregate into discrete morphophysiological or geographic clusters. However, evolved cultivar-groups (cv.-gr. Melanophthalmus and cv.-gr. Sesquipedalis) appear to be less diverse than their putative primitive cultivar-group progenitors. Due to the lack of availability of critical material, no clear center of origin can be established. However, the data presented suggest that Northeast Africa could be a possible center of domestication.

An electrophoretic comparison of variation at 29 isozyme loci was performed for 21 Vigna accessions belonging to V. frutescens , V. membranacea and V. friesiorum . The results do not fit accepted nomenclature and the taxonomic ranks should be reassessed. V. frutescens , V. friesiorum , and various subspecies of V. membranacea should be considered as putative species, although hybridisation data are needed in order to obtain a clear picture of the organisation of this gene pool. However, the results do not support the existence of two sections, and section Liebrechtsia is merged with section Macrodontae in a newly enlarged section Macrodontae.

The cowpea [Vigna unguiculata (L.) Walp.] is a morphologically and genetically variable species composed of wild perennial, wild annual, and cultivated forms that are mainly used for edible seeds and pods. In this study, genetic variation in 199 germplasm accessions of wild and cultivated cowpea was evaluated using an allozyme analysis. The results from this survey showed that wild cowpea exhibits genetic variation perfectly fitted with the existing morphological classification. The cowpea gene-pool is characterized by its unusually large size. It encompasses taxa (ranked as subspecies) that could be considered as different species considering the high genetic distances observed between accessions belonging to different taxa. These subspecies can be classified into three groups characterized by their breeding systems: perennial outcrossers, perennial out-inbreds, and inbred annuals. Allozyme data confirm this grouping. Perennial outcrossers look primitive and are more remote from each other and from perennial out-inbreds. Within this large gene-pool, mainly made of perennial taxa, cultivated cowpeas (ssp. unguiculata var. unguiculata) form a genetically coherent group and are closely related to annual cowpeas (ssp. unguiculata var. spontanea) which may include the most likely progenitor of cultivated cowpeas.

An electrophoretic comparison of variation at 28 isozyme loci was performed for 58Vigna accessions belonging to theV. luteola, V. ambacensis, andV. racemosa groups of species. In all three groups, strong divergence is noted between results and actual nomenclature.

Background Cowpea is a highly inbred crop. It is part of a crop-weed complex, whose origin and dynamics is unknown, which is distributed across the African continent. This study examined outcrossing rates and genetic structures in 35 wild cowpea (Vigna unguiculata ssp. unguiculata var. spontanea) populations from West Africa, using 21 isozyme loci, 9 of them showing polymorphism. Results Outcrossing rates ranged from 1% to 9.5% (mean 3.4%), which classifies the wild cowpea breeding system as primarily selfing, though rare outcrossing events were detected in each population studied. Furthermore, the analyses of both the genetic structure of populations and the relationships between the wild and domesticated groups suggest possibilities of gene flow that are corroborated by field observations. Conclusions As expected in a predominantly inbred breeding system, wild cowpea shows high levels of genetic differentiation and low levels of genetic diversity within populations. Gene flow from domesticated to wild cowpea does occur, although the lack of strong genetic swamping and modified seed morphology in the wild populations suggest that these introgressions should be rare.

International audience; SymRK is one of the key genes involved in initial steps of legume symbiotic association with fungi (mycorrhization) and nitrogen-fixing bacteria (nodulation). A large portion of the sequence encoding the extracellular domain of SYMRK was obtained for 38 lupine accessions and 2 outgroups in order to characterize this region, to evaluate its phylogenetic utility, and to examine whether its molecular evolutionary pattern is correlated with rhizobial diversity and specificity in Lupinus. The data suggested that, in Lupinus, SymRK is a single copy gene that shows good phylogenetic potential. Accordingly, SymRK provided additional support to previous molecular phylogenies, and shed additional light on relationships within the Old World group of Lupinus, especially among the African species. Similar to results of other studies, analyses of SymRK sequences were unable to resolve placement of the Florida unifoliolate lineage, whose relationship was weakly supported to either the Old or the New World lupines. Our data are consistent with strong purifying selection operating on SymRK in Lupinus, preserving rather than diversifying its function. Thus, although SymRK was demonstrated to be a vital gene in the early stages of the root-bacterial symbiotic associations, no evidence from present analyses indicate that this gene is involved in changes in rhizobial specificity in Lupinus.

Amplified fragment length polymorphisms (AFLPs) were used to evaluate genetic relationships within cowpea (Vigna unguiculata (L.) Walp.) and to assess the organization of its genetic diversity. Nei's genetic distances were estimated for a total of 117 acces- sions including 47 domesticated cowpea (ssp. ungui- culata var. unguiculata), 52 wild and weedy annuals (ssp. unguiculata var. spontanea), as well as 18 perennial accessions of the wild subspecies pubescens, tenuis and alba. AFLP variation was also used to study genetic variation among and within domesticated and wild ac- cessions based on their geographical origin (western, eastern and southern Africa). Wild annual cowpea (var. spontanea) (H T =0.175) was more diverse than domesti- cated cowpea (H T =0.108). Wild cowpea was more di- verse in eastern (HS=0.168) than in western Africa (H S =0.129), suggesting an eastern African origin for the wild taxon. The AFLP data were consistent with earlier findings of a unique domestication event in cowpea in the northern part of the continent and suggested that do- mestication in eastern or southern Africa was unlikely. It did not allow a more precise localization of domestica- tion due to extensive gene flow between wild and do- mesticated forms that has led to a large crop-weed com- plex distributed over the entire African continent. In ad- dition, wild materials from northeastern Africa are still lacking. Overall, the superiority of the AFLP technique over isozymes resided in its ability to uncover variation both within domesticated and wild cowpea, and should be a powerful tool once additional wild material be- comes available.

Bambara groundnut (#Vigna subterranea$ (L.) Verdc.) is an important crop in many countries of tropical Africa. Compared with other leguminous crops, little is known about the organization of its genetic diversity. Our goal was to investigate its population structure and the partitioning of genetic diversity between domesticated and wild accessions. Seventy-nine accessions of domesticated and 21 wild Bambara groundnut populations were evaluated for genetic diversity at 41 isozyme loci, representing 23 enzyme systems. Domesticated accessions were characterized by very low genetic diversity (Ht = 0.052) with only 7 polymorphic loci. Wild populations were characterized by higher genetic diversity (Ht = 0.087), with 14 polymorphic. This suggests a marked bottleneck between wild and domesticated forms. Intrapopulation diversity was comparatively high (Hs = 0.033 for domesticated and Hs = 0.025 for wild populations) despite the near absence of heterozygous individuals, which suggests a predominantly selfing mode of pollination in both wild and domesticated Bambara groundnut. High genetic identity between wild and domesticated forms suggests that wild Bambara groundnut is the true progenitor of domesticated Bambara groundnut. Both morphological and isozyme data show a gradient between true wild and domesticated Bambara groundnut through weedy populations. The improved understanding of the organization of genetic diversity of Bambara groundnut provided by our data will allow a better conservation of its genetic resources. (Résumé d'auteur)