Haby Sanou

Genetic diversity of the shea tree (Vitellaria paradoxa C.F. Gaertn), detected by RAPD and chloroplast microsatellite markers.

RAPDs and chloroplast microsatellites were used to quantify the genetic variation of Vitellaria paradoxa (an economically important tree species in sub-Saharan Africa, north of the equator) and to analyse the geographic distribution of diversity in relation to the refuge theory. A total of 13 locations were sampled in eight countries, covering most of the natural range from Senegal to Uganda. A total of 67 polymorphic and 15 monomorphic RAPD loci were detected in 179 individuals. No relationship was identified between diversity and longitude or latitude. An unrooted neighbour-joining tree suggested a western group and an eastern group, representing 7% (P=0.000) of the total variation. A Mantel test suggested that genetic distances between populations were correlated to geographic distances (R=0.88, P=0.001). The three-chloroplast microsatellite primers, assayed in 116 individuals, revealed 10 different alleles and seven chlorotypes. Most of the populations comprised a single haplotype. It is proposed from these results that the difference between western and eastern populations results from putative refugia separated by the current ‘Dahomey Gap’ (a semiarid zone that meets the coast around the Ghana–Togo–Benin–Nigeria borders), an area that may have been exceptionally dry during glacial periods. In addition, it is suggested that the haplotype distribution and frequency in the western populations could be due to the more recent impact of humans, particularly shea tree selection and dispersal during traditional agroforestry.

Comparison of quantitative and molecular variation in agroforestry populations of the shea tree (Vitellaria paradoxa C.F. Gaertn) in Mali

In this study we investigated the within‐ and between‐population genetic variation using microsatellite markers and quantitative traits of the shea tree, Vitellaria paradoxa, an important agroforestry tree species of the Sudano–Sahelian region in Africa. Eleven populations were sampled across Mali and in northern Côte d’Ivoire. Leaf size and form and growth traits were measured in a progeny test at the nursery stage. Eight microsatellites were used to assess neutral genetic variation. Low levels of heterozygosity were recorded (1.6–3.0 alleles/locus; HE = 0.25–0.42) and the fixation index (FIS = −0.227–0.186) was not significantly different from zero suggesting that Hardy–Weinberg equilibrium is encountered in all populations sampled. Quantitative traits exhibited a strong genetic variation between populations and between families within populations. The degree of population differentiation of the quantitative traits (QST = 0.055–0.283, QSTmean = 0.189) strongly exceeds that in eight microsatellite loci (FST = −0.011–0.142, FSTmean = 0.047). Global and pairwise FST values were very low and not significantly different from zero suggesting agroforestry practices are amplifying gene flow (Nm = 5.07). The population means for quantitative traits and the rainfall variable were not correlated, showing variation was not linked with this climatic cline. It is suggested that this marked differentiation for quantitative traits, independent of environmental clines and despite a high gene flow, is a result of local adaptation and human selection of shea trees. This process has induced high linkage disequilibrium between underlying loci of polygenic characters.

Advances in domestication of indigenous fruit trees in the West African Sahel

Fruit trees play an important nutritional role for livelihoods of rural people in the West African Sahel through provision of energy and nutrients such as vitamins, minerals, and proteins. Research on the domestication of local fruit trees has started recently through projects concentrating on some of the most important indigenous species of dry West Africa, i.e. Adansonia digitata, Parkia biglobosa, Tamarindus indica, Vitellaria paradoxa and Ziziphus mauritiana. We present a status of finalised and ongoing domestication research with the aim of defining research gaps that would need to be covered by future research activities to obtain higher yields and better quality fruits. Germplasm collection in central West Africa has been intense compared to elsewhere in the species’ distribution areas, but conservation status of the material is poor since it is only planted in few trials. Knowledge of genetic parameters, especially for fruit traits, is almost absent, but characterisation of genotypes is underway for some of the species. Mating systems and patterns are still unknown for many species. Efficient vegetative propagation based on simple techniques was shown to be possible for all species except P. biglobosa. In order to secure immediate as well as long term gains, we recommend combining clonal propagation of selected plus individuals with recombination and breeding of selected genotypes. We discuss whether local institutions in the Sahel have the financial capacity to carry out long term breeding programmes, and suggest that efforts should be made to find new ways of disseminating improved germplasm.

Phenotypic variation of agromorphological traits of the shea tree, Vitellaria paradoxa C.F. Gaertn., in Mali

Forty-one shea tree populations were sampled, spanning the main climatic zones of Vitellaria paradoxa Gaertn. in Mali and 10–35 adult trees were chosen randomly in the agroforestry parklands of each area. A total of 12 morphological traits, related to tree morphology, fruit size and leaf form were measured. The variance components showed that variation among populations represented the smaller percentage of the total variation with most of the values varying between 15 and 30%. The repeatability coefficient was generally high for tree within populations with values ranging between 0.23 and 0.78. Although genetic correlations cannot be accurately estimated, due to difficulties in separation from environmental effects, the results indicate that there is a very low genetic relation between the three kinds of traits, i.e., between those related to tree, those related to leaf and those related to fruit. Leaf and fruit size traits were positively and significantly correlated with rainfall, although tree circumference was negatively correlated with rainfall and the significantly larger shea trees were noted in the drier areas – an observation thought linked to human management of the parklands. Soil drainage and parkland density, however, did not explain differences between populations for fruit traits. This study offers preliminary information for the development of a breeding population for a shea tree improvement programme. The value of repeatability, the low correlation between sets of traits and the distribution of variation, suggest that selection of many individual trees within a few populations, would allow capture of large genetic gain especially for fruit traits.

An analysis of the pattern of genetic variation in Vitellaria paradoxa using RAPD markers

Vitellaria paradoxa C.F.Gaertn. is one of the most economically and socially important tree species in the Sudano-Sahelian region. Little is known of the pattern of variation within its natural range. Eight populations covering most of the natural range from Senegal to Uganda were sampled and leaves of 118 individual trees were collected. An analysis of molecular diversity was carried out using random amplified polymorphic DNA (RAPD) markers. Fifteen random primers generated 67 polymorphic and 15 monomorphic RAPD loci ranging from size 1670 bp to 280 bp. Shannons diversity index varied from Central Africa/Ndele (0.374) to Uganda/Amoya (0.350) but the differences between populations were smaller than the population standard errors. Correspondence analysis of unrooted neighbour-joining trees suggested that genetic distances between populations were correlated with geographic distances. This trend was confirmed by a Mantel test giving a coefficient of correlation between genetic and geographic distances of R = 0.88 (P = 0.0001). Result of AMOVA (analyses of molecular variance) showed that 14.8% (P = 0.002) of the RAPD variation was distributed among populations. Nested analysis of variance indicated that variance between the western and eastern groups of population represented 8.7% (P = 0.001) of the total variation and the variation amongst populations within group was 9.5% (P = 0.001). Eighty two percent of the variation was explained by variation amongst individuals within populations. The origin of genetic structure and level of diversity may be explained by the glacial refugia, the biological traits of Vitellaria paradoxa and by the impact of semi-domestication. Based on these results, sampling options of the natural populations are suggested for in or ex situ conservation. For the development of Vitellaria paradoxa breeding population, the sampling should consist of many individual trees selected within a few populations to capture a large proportion of variation.

A research approach supporting domestication of Baobab (Adansonia digitata L.) in West Africa

The Baobab tree (Adansonia digitata L.) is a key multipurpose species for the African region. In the recent years there has been an extended commercial interest for different A. digitata products. As a spectacular African key species there has been a growing interest from NGO′s and various research groups. A research group, focussing on the following countries Burkina Faso, Mali and Niger, has participated in a concerted research action with cross counter disciplinary cooperation between plant physiology, population genetics, tree breeding, food science, and socioeconomics. This paper presents a review and the way knowledge gaps are being addressed using the above mentioned approach. The overall work was initiated in 2005–2006 when a large collection of A. digitata seeds was carried out in 15 African countries. Fourteen populations were selected in Burkina Faso, Mali, and Niger including more than 400 superior trees, and many of the trees have been selected along with farmers in a participatory process. Studies have been initiated in all countries on geographic variation of growth, adaptive and phenological traits. This includes studies in the nursery, establishment of international provenance series, seed orchards and studies of gene flow and phylogeographic variation with various markers. Specific nursery trials have been established in order to study provenance and progeny performance related to drought stress. Several stress related characteristics are being measured. Preliminary observations showed large morphological variation between African provenances of A. digitata. Food properties of plant parts have been studied within and between populations. This includes studies of total biomass production, vitamin A, B1, B2, and various minerals. The influence of stress will be related to these parameters. Fruits, which are rich in sugar and vitamin C, are used as an ingredient in juice and other foods. Seeds can be used directly as food ingredient or in fermented condition (Maari). The fermentation process was previously poorly described. Therefore, the microorganisms associated with Baobab seeds fermentation have been identified and a starter cultures for control production has been proposed. The efficiency of dissemination of superior A. digitata trees depends on vegetative propagation. Various grafting methods are currently being tested in cooperation with farmers. Micropropagation is tested as well as a method for clone propagation. The accumulated knowledge will be applied for a domestication strategy of A. digitata in Burkina Faso, Mali and Niger. The outcome from this study will be guidelines for protection and management of the germplasm of A. digitata resources.

Molecular data reveal isolation by distance and past population expansion for the shea tree (Vitellaria paradoxa C.F. Gaertn) in West Africa.

While the genetic structure of many tree species in temperate, American and Asian regions is largely explained by climatic oscillations and subsequent habitat contractions and expansions, little is known about Africa. We investigated the genetic diversity and structure of shea tree (Vitellaria paradoxa,) in Western Africa, an economically important tree species in the Sudano‐Sahelian zone. Eleven nuclear microsatellites (nuc) were used to genotype 673 trees selected in 38 populations. They revealed moderate to high within‐population diversity: allelic richness ranged from Rnuc = 3.99 to 5.63. This diversity was evenly distributed across West Africa. Populations were weakly differentiated (FSTnuc = 0.085; P < 0.0001) and a pattern of isolation by distance was noted. No phylogeographic signal could be detected across the studied sample. Additionally, two chloroplast microsatellite loci, leading to 11 chlorotypes, were used to analyse a sub‐set of 370 individuals. Some variation in chloroplast allelic richness among populations could be detected (Rcp = 0.00 to 4.36), but these differences were not significant. No trend with latitude and longitude were observed. Differentiation was marked (GSTcp = 0.553; P < 0.0001), but without a significant phylogeographical signal. Population expansion was detected considering the total population using approximate Bayesian computation (nuclear microsatellites) and mismatch distribution (chloroplast microsatellites) methods. This expansion signal and the isolation by distance pattern could be linked to the past climatic conditions in West Africa during the Pleistocene and Holocene which should have been favourable to shea tree development. In addition, human activities through agroforestry and domestication (started 10 000 bp) have probably enhanced gene flow and population expansion.

Isolation and characterization of new microsatellite markers in shea tree (Vitellaria paradoxa C. F. Gaertn)

Vitellaria paradoxa is one of the major components of African parkland agroforestry systems. In order to assess the genetic diversity and population structure of this species, we isolated and characterized 14 polymorphic nuclear microsatellite loci. Primers developed to amplify these loci were used to analyse 200 individuals of a shea tree population in Mali. Loci have shown a high number of alleles ranging from four to 26, and display an observed level of heterozygosity between 0.37 and 0.85. These new very polymorphic microsatellite markers will be useful for genetic and ecological studies of V. paradoxa.

Breeding for high production of leaves of baobab (Adansonia digitata L) in an irrigated hedge system

This study presents an evaluation of an irrigated “market garden” trial established with 3-month-old seedlings of baobab. The trial included offspring from 59 open-pollinated family lots from ten provenances and four bulked provenance sample lots. Leaf productivity and seedling growth were evaluated monthly from the third month after establishment. We found lowered leaf productivity during the dry season despite the plants being irrigated. We provide the first estimates of heritability for leaf production and growth of the species. We assumed that the families of seed from open-pollinated single trees consist of true half-sibs but also provided adjusted estimates assuming a high level of selfing. Differences among provenances and families within provenances were highly significant (p < 0.01) with respect to leaf productivity. Heritability estimates (adjusted and non-adjusted) for leaf productivity were moderate to low, predicting moderate genetic gain from selection. Strong genetic and phenotypic correlations were estimated between diameter and leaf production, indicating that simple selection for diameter can efficiently increase leaf production. The dry weight/fresh weight ratio was not significantly different among provenances or families within provenances. Based on the findings, we discuss how breeding can increase the total leaf production and its seasonal distribution.

Yield and biomass production by African eggplant (Solanum aethiopicum) and sorghum (Sorghum bicolor) intercropped with planted Ber (Ziziphus mauritiana) in Mali (West Africa)

Methods for introduction of an improved cultivar of Ber tree (Ziziphus mauritiana Lam.) also called Jujube and its management on farms through intercropping with African eggplant (Solanum aethiopicum L.) and sorghum [Sorghum bicolor (L.) Moench] were tested in Sanankoroba, Mali. An experimental plantation was established on-farm with seedlings of the local variety of Ber half of which were grafted in situ with an improved cultivar of Ber called Seb. Crop production, Ber growth and fruit production were assessed over two cropping seasons. Both the improved and local varieties of Ber had no detrimental effect on either eggplant or sorghum, both in terms of yield and nutritional quality. In fact a beneficial effect of trees was found on the performance of both crops (yield and dry matter production) which suggests complementarities in resource use. The high level of fruit production by the improved variety of Ber observed on farms under rain-fed conditions may be a source of additional income and diversification of diet for rural communities in West Africa. Therefore, farmers’ adoption of growing improved Ber varieties in association with food crops may help considerably in improving food security and alleviating poverty in the region.