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Bambara groundnut has the potential to be used to contribute more the climate change ready agriculture. The requirement for nitrogen fixing, stress tolerant legumes is clear, particularly in low input agriculture. However, ensuring that existing negative traits are tackled and demand is stimulated through the development of markets and products still represents a challenge to making greater use of this legume. World agriculture is currently based on very limited numbers of crops, representing a significant risk to food supplies, particularly in the face of climate change which is expected to increase the frequency of extreme events. Minor and underutilised crops can help to develop a more resilient and nutritionally dense future agriculture. Bambara groundnut [Vigna subterranea (L.) Verdc.[, as a drought resistant, nitrogen-fixing, legume has a role to play. However, as with most underutilised crops, there are significant gaps in knowledge and also negative traits such as 'hard-to-cook' and 'photoperiod sensitivity to pod filling' associated with the crop which future breeding programmes and processing methods need to tackle, to allow it to make a significant contribution to the well-being of future generations. The current review assesses these factors and also considers what are the next steps towards realising the potential of this crop.

© Copyright © 2020 Tan, Azam-Ali, Goh, Mustafa, Chai, Ho, Mayes, Mabhaudhi, Azam-Ali and Massawe. Rapid population growth, climate change, intensive monoculture farming, and resource depletion are among the challenges that threaten the increasingly vulnerable global agri-food system. Heavy reliance on a few major crops is also linked to a monotonous diet, poor dietary habits, and micronutrient deficiencies, which are often associated with diet-related diseases. Diversification—of both agricultural production systems and diet—is a practical and sustainable approach to address these challenges and to improve global food and nutritional security. This strategy is aligned with the recommendations from the EAT-Lancet report, which highlighted the urgent need for increased consumption of plant-based foods to sustain population and planetary health. Bambara groundnut (Vigna subterranea (L.) Verdc.), an underutilized African legume, has the potential to contribute to improved food and nutrition security, while providing solutions for environmental sustainability and equity in food availability and affordability. This paper discusses the potential role of Bambara groundnut in diversifying agri-food systems and contributing to enhanced dietary and planetary sustainability, with emphasis on areas that span the value chain: from genetics, agroecology, nutrition, processing, and utilization, through to its socioeconomic potential. Bambara groundnut is a sustainable, low-cost source of complex carbohydrates, plant-based protein, unsaturated fatty acids, and essential minerals (magnesium, iron, zinc, and potassium), especially for those living in arid and semi-arid regions. As a legume, Bambara groundnut fixes atmospheric nitrogen to improve soil fertility. It is resilient to adverse environmental conditions and can yield on poor soil. Despite its impressive nutritional and agroecological profile, the potential of Bambara groundnut in improving the global food system is undermined by several factors, including resource limitation, knowledge gap, social stigma, and lack of policy incentives. Multiple research efforts to address these hurdles have led to a more promising outlook for Bambara groundnut; however, there is an urgent need to continue research to realize its full potential.

In Bambara groundnut [Vigna subterranea (L) Verdc], long photoperiods can cause a decline in pod and seed number. While negative photoperiod effects on pod yield have been reported, positive effects and the existence of genotypes less sensitive for photoperiod have not been reported. Ten geographically diverse genotypes were evaluated over 3 years under short and long photoperiod for yield‐related traits (pod number, pod weight, seed number, seed weight, and 100 seed weight), pod growth habit (geocarpic or ageocarpic pods), peduncle elongation and soil penetration, and pod set/abortion. Anatomical sections of gynophores and embryonic pod development of a highly photoperiod sensitive genotype “Ankpa‐4” from Nigeria were examined. A strong photoperiodic effect on yield component traits (pod number, pod weight, seed number, seed weight, and 100 seed weight) was observed and with considerable variation between genotypes. Three classes of photoperiod effect on pod yield were identified, (a) qualitative short‐day types; (b) quantitative short‐day types; and (c) quantitative long‐day types. In long photoperiods, above‐ground vegetative biomass and the length of lateral branches in some genotypes increased by at least twofold. Morphological and anatomical characterization of gynophores and developing pods of the most sensitive line shows continuous geocarpic growth, but with healthy embryonic pods failing to enlarge after soil penetration. Results from the yield patterns of the three phenotypic classes confirm that pods and seeds are high priority sinks, and long photoperiod alters the balance in assimilate distribution between competing yield forming processes and vegetative sinks. By exploiting the genetic variation characterized here for photoperiod requirement for pod filling, the geographical range of this crop could be extended beyond current growing regions. In the near future, we anticipate that Bambara groundnut will become a significant contributor to global food, nutritional, and environmental security once these photoperiod issues are resolved.

Main conclusion\ud \ud Crops For the Future (CFF), as an entity, has established a broad range of research activities to promote the improvement and adoption of currently underutilised crops.\ud \ud \ud Abstract\ud \ud This paper summarises selected research activities at Crops For the Future (CFF) in pursuit of its mission ‘to develop solutions for diversifying future agriculture using underutilised crops’. CFF is a research company focussed on the improvement of underutilised crops, so that they might be grown and consumed more widely with benefits to human food and nutritional security; its founding guarantors were the Government of Malaysia and the University of Nottingham. From its base in Malaysia, it engages in research around the world with a focus on species and system diversification. CFF has adopted a food system approach that adds value by delivering prototype food, feed and knowledge products. Bambara groundnut (Vigna subterranea) was adopted as an exemplar crop around which to develop CFF’s food system approach with emphasis on the short-day photoperiod requirement for pod-filling and the hard-to-cook trait. Selective breeding has allowed the development of lines that are less susceptible to photoperiod but also provided a range of tools and approaches that are now being exploited in other crops such as winged bean (Psophocarpus tetragonolobus), amaranth (Amaranthus spp.), moringa (Moringa oleifera) and proso (Panicum miliaceum) and foxtail (Setaria italica) millets. CFF has developed and tested new food products and demonstrated that several crops can be used as feed for black soldier fly which can, in turn, be used to feed fish thereby reducing the need for fishmeal. Information about underutilised crops is widely dispersed; so, a major effort has been made to develop a knowledge base that can be interrogated and used to answer practical questions about potential exploitation of plant and nutritional characteristics. Future research will build on the success with Bambara groundnut and include topics such as urban agriculture, rural development and diversification, and the development of novel foods.

SUMMARYThe effect of drought and temperature on the growth and development of bambara groundnut (Vigna subterranea (L.) Verdc.) was studied in controlled environment glasshouses in the United Kingdom. There were two landraces, S19-3 (from a hot, dry environment; Namibia) and Uniswa Red (from a cool, wet environment; Swaziland), two temperature regimes (23 °C and 33 °C) and three watering regimes (2006; fully irrigated), 2007 (drought imposed at 77 days after sowing (DAS)) and 2008 (drought imposed at 30 DAS)). Bambara groundnut responded to drought by slowing the rate of leaf area expansion and reducing final canopy size and total dry matter (TDM). Drought also caused significant reductions in pod dry matter, pod numbers and harvest index (HI), leading to a decrease in final yield that was different between landraces. Throughout the three growing seasons, landraces grown at 33 °C produced more TDM than the landraces grown at 23 °C. The two landraces differed in their phenology; S19-3 exhibited a reduced phenology where leaf numbers started to decrease before Uniswa Red at both temperatures, while Uniswa Red maintained the longest life cycle. The lowest pod yield was produced by Uniswa Red in 2008 at 33 °C (maximum of 35.5 gm−2), while S19-3 produced a minimum pod yield of 56.6 gm−2 at 33 °C, also in 2008. However, both landraces produced considerably more pod yield at 23 °C throughout the three growing seasons (minimum of 151 gm−2 and 162 gm−2 for Uniswa Red and S19-3, respectively). Under moderate drought, S19-3 at 33 °C gave the highest pod yield (365 gm−2) among the treatments throughout the three growing seasons and maintained HI better under drought. Despite being from a hot, dry environment, S19-3 also performed well at low temperature, which indicates the adaptation of S19-3 to low temperature that it also experiences in the country of origin.

SUMMARYSimulation of yield response to water plays an increasingly important role in optimization of crop water productivity (WP) especially in prevalent drought in Africa. The present study is focused on a representative crop: bambara groundnut (Vigna subterranea), an ancient grain legume grown, cooked, processed and traded mainly by subsistence women farmers in sub-Saharan Africa. Over four years (2002, 2006–2008), glasshouse experiments were conducted at the Tropical Crops Research Unit, University of Nottingham, UK under controlled environments with different landraces, temperatures (23 ± 5 °C, 28 ± 5 °C, 33 ± 5 °C) and soil moisture regimes (irrigated, early drought, late drought). Parallel to this, field experiments were conducted in Swaziland (2002/2003) and Botswana (2007/2008). Crop measurements of canopy cover (CC), biomass (B) and pod yield (Y) of selected experiments from glasshouse (2006 and 2007) and field (Botswana) were used to calibrate the FAO AquaCrop model. Subsequently, the model was validated against independent data sets from glasshouse (2002 and 2008) and field (Swaziland) for different landraces. AquaCrop simulations for CC, B and Y of different bambara groundnut landraces are in good agreement with observed data with R2 (CC-0.88; B-0.78; Y-0.72), but with significant underestimation for some landraces.

Genetic diversity of a Bambara groundnut germplasm representing accessions from 25 African countries, maintained at the International Institute of Tropical Agriculture (IITA; Nigeria) was evaluated based on seed patterns, qualitative characters, quantitative traits and Diversity Arrays Technique (DArT) markers. The study aimed at identifying important descriptors for germplasm conservation, validating the crop’s geographical centre of origin and facilitating the utilization of existing genetic resources. Frequencies and diversity indices of seven types of seed patterns, 12 qualitative and 28 quantitative traits were evaluated for 124 representative accessions and were analyzed by region to elucidate the geographic distribution of descriptor states. In addition, individual plant samples from a subset of 40 landrace accessions were analyzed by assessing 554 DArT markers. Both the phenetic tree constructed from the qualitative and quantitative descriptors and the population structure derived from DArT marker analysis suggested a relatively high genetic diversity among accessions. Higher genetic diversity was observed for the Cameroon/Nigeria region relative to other regions, in agreement with the hypothesis that this region is the centre of origin for Bambara groundnut. Use of an extensive and diverse range of germplasm and an approach that integrated morphological and quantitative descriptors with DNA markers that represent wide genomic coverage offered a powerful way to make inferences about crop germplasm, in support of crop-improvement programs.

Canopy development of bambara groundnut (Vigna subterranea (L.) Verdc) is affected by temperature stress, drought stress and photoperiod. The quantification of these documented effects by means of a suitable crop model, BAMGRO is presented in this paper. Data on canopy development from five growth chamber, four glasshouse and three field experiments were analyzed to calibrate and validate the BAMGRO model to produce simulations for temperature stress, drought stress and photoperiodic effect on two contrasting landraces; Uniswa Red (Swaziland) and S19-3 (Namibia). The daily initiation rate of new leaves is calculated by means of a Gaussian function and is altered by temperature stress, drought stress, photoperiod and plant density. The rate in dead leaf number is dependent upon the maximum senescence fraction which can be explained by physiological maturity, mutual shading, temperature stress and drought stress. The leaf area expansion is calculated as a function of leaf number and individual leaf size. The canopy development of bambara groundnut is modelled (and tested) in BAMGRO model by means of leaf initiation and leaf area expansion and branching and stem production are not considered. Nash and Sutcliffe value (N–S) is used as the measure of the mean square error to the observed variance. The simulated leaf number per plant correlated well with the glasshouse measurements for Uniswa Red for the two considered years (2007 and 2008) with high N–S value (Nash and Sutcliffe; from 0.76 to 0.92), low mean absolute error (MAE; from ±2.91 to 3.98) and Botswana field grown in January 18 and February 1, 2007 (N–S, 0.88 and 0.74; MAE, ±0.59 and ±6.18). Overall, simulated LAI correlated well with measured values for the two tested landraces; S19-3 (N–S value, 0.84) and Uniswa Red (N–S value, 0.80) with maximum MAE less than ±0.50 thus explaining canopy development under temperature stress, drought stress and variable photoperiods.

The effect of drought on the growth and development of bambara groundnut (Vigna subterranea (L.) Verdc.) was studied in controlled-environment glasshouses in the UK. There were three landraces (S19-3, DipC and UN from Namibia, Botswana and Swaziland, respectively) and two watering regimes; a control that was irrigated weekly to 90% field capacity and a drought treatment with no irrigation from 49 days after sowing (DAS) until final harvest (147 DAS). Bambara groundnut responded to drought by reducing the rate of leaf area expansion, final canopy size and total dry matter (TDM) during vegetative growth. Drought also caused significant reductions in pod dry matter (PDM), pod number, seed weight and harvest index (HI), leading to a decrease in final pod yield that was different between landraces. Across landraces, drought reduced mean pod yield from 298 g m−2 to 165 g m−2, representing 45% yield loss. Despite the reduction in all landraces, the mean pod yield across the droughted treatments that had received no water for almost 100 days indicated the resilience of the species to drought. The three landraces differed in their phenology; S19-3 exhibited a reduced phenology while UN maintained the longest life cycle. The different responses of the landraces reflect their adaptation to their local climates where mean annual rainfall ranges between 365 mm (Namibia) and 1390 mm (Swaziland). We discuss the significance of these results for future breeding programmes on bambara groundnut.

Controlled crosses in bambara groundnut were attempted between a range of thirty-six bambara groundnut landraces (thirty domesticated (V. subterranea var. subterranea) and six wild (V. subterranea var. spontanea)). Ten F-1 seed were produced. Of these, eight germinated producing F-2 populations. On seed set, four populations could be unambiguously confirmed as true crosses by F-3 seed coat colour. A single F-2 population, derived from a domesticated landrace from Botswana (DipC; female parent) crossed with a wild accession collected in Cameroon (VSSP11; male parent) was used to study a range of agronomic and domestication traits. These included; days to emergence, days to flowering, internode (fourth) length at harvest, number of stems per plant, leaf area, Specific Leaf Area (SLA), Carbon Isotope Discrimination (CID), 100 seed weight, testa colour and eye pattern around the hilum. On the basis of variation for internode length and stems per plant, 14 small F-3 families were selected and grown under field conditions to further investigate the genetic basis of the 'spreading' versus 'bunched' plant character, a major difference between wild and cultivated bambara groundnut. Results presented suggest that traits including leaf area, SLA, CID and 100 seed weight are controlled by several genes. In contrast, the variation for traits such as internode length, stems per plant, days to emergence and seed eye pattern around the hilum are likely to be under largely monogenic control. The results of this work are discussed in relation to the domestication of bambara groundnut.

Ten bambara groundnut [Vigna subterranea (L.) Verdc] landraces of diverse origin were used to examine the effect of temperature on the rate of leaf appearance (RLA) and to determine the base temperature (T b ) among landraces. Knowledge of leaf development is required in breeding programs where crop morphological development is considered an important selection criterion. The ability to predict leaf appearance would also facilitate modeling of plant development. Plants were grown in controlled-environment growth rooms under different temperatures (20, 25, 30, 32, and 35°C ± 1). The study to examine the effect of temperature on leaf appearance in bambara groundnut landraces indicated that the base temperature (T b ) for leaf appearance ranged from 8.1 to 12.0°C. Within the range of temperatures evaluated in this study, the observation of differences in T b values among the landraces suggests that T b values for leaf appearance are landrace dependent. The phyllochron was also variable among landraces ranging from 40.9 to 53.0°C. The RLA ranged from 0.19 to 0.63 leaves d -1 depending on temperature and landrace. The landrace variations in relation to leaf appearance may allow selection of landraces for different environments.

Genetic diversity in 12 landraces of bambara groundnut (Vigna subterranea), an indigenous African legume, was evaluated using Random Amplified Polymorphic DNA (RAPD) markers. DNA from individuals of each landrace was also analysed to determine the level of heterogeneity within landraces. RAPDs revealed high levels of polymorphism among landraces. The percentage polymorphism ranged from 63.2% to 88.2% with an average of 73.1% for the 16 RAPD primers evaluated. The construction of genetic relationships using cluster analysis groups the 12 landraces in two clusters. RAPDs are useful for the genetic diversity studies in V. subterranea and can identify variation within landraces.

Bambara groundnut (Vigna subterranea (L.) Verdc), an African indigenous legume, is popular in most parts of Africa. The present study was undertaken to establish genetic relationships among 16 cultivated bambara groundnut landraces using fluorescence-based amplified fragment length polymorphism (AFLP) markers. Seven selective primer combinations generated 504 amplification products, ranging from 50 to 400 bp. Several landrace-specific products were identified that could be effectively used to produce landrace-specific markers for identification purposes. On average, each primer combination generated 72 amplified products that were detectable by an ABI Prism 310 DNA sequencer. The polymorphisms obtained ranged from 68.0 to 98.0%, with an average of 84.0%. The primer pairs M-ACA + P-GCC and M-ACA + P-GGA produced more polymorphic fragments than any other primer pairs and were better at differentiating landraces. The dendrogram generated by the UPGMA (unweighted pair-group method with arithmetic averaging) grouped 16 landraces into 3 clusters, mainly according to their place of collection or geographic origin. DipC1995 and Malawi5 were the most genetically related landraces. AFLP analysis provided sufficient polymorphism to determine the amount of genetic diversity and to establish genetic relationships in bambara groundnut landraces. The results will help in the formulation of marker-assisted breeding in bambara groundnut.Key words: under-utilized, African legume, molecular markers.

Three landraces of bambara groundnut (Vigna subterranea (L.) Verdc.) were grown as crop stands in controlled environment glasshouses at the Tropical Crops Research Unit, University of Nottingham, in 1995. Two soil moisture treatments were imposed: irrigated to 90% field capacity each week and irrigated to 60% field capacity until establishment (27 days after sowing) with no further irrigation. Seasonal mean fractional interception varied between 0·20–0·37 for the droughted treatments and 0·62–0·74 for the irrigated treatments, resulting in cumulative intercepted radiation of 228–350 MJ/m2 and 662–794 MJ/m2, respectively. The maximum total dry matter (DM) produced was 5·8 t/ha at final harvest (145 days after sowing) with a pod yield of 2·7 t/ha. Under moisture stress there was little difference in DM production between landraces, with the highest total DM of 1·1 t/ha and a pod yield of 0·05 t/ha, representing a harvest index of 0·05 compared with an average of 0·46 for the irrigated treatments. The conversion coefficient was reduced from 1·00 under irrigation to 0·51 g DM/MJ radiation intercepted by soil moisture stress. Two of the landraces showed adaptive mechanisms to avoid drought; these are discussed in relation to maximizing seasonal radiation interception.

Bambara groundnut is an indigenous African legume crop plant. It is largely grown by subsistence farmers, but can also be used as a cash crop to supplement family income. The fact that it is highly drought tolerant means that it has considerably potential to provide part of food security in regions of the world where water availability is a serious issue. As one part of the international effort to understand and improve this species, we report here the first 10 microsatellite markers derived from bambara groundnut.

The components of leaf water potential (Ψ I ) and relative water content (RWC) were measured for stands of bambara groundnut (Vigna subterranea) exposed to three soil moisture regimes in controlled-environment glasshouses at the Tropical Crops Research Unit, Sutton Bonington Campus. Treatments ranged from fully irrigated (wet) to no irrigation from 35 days after sowing (DAS) (dry). RWC values varied between 92-96% for the wet treatment, but declined from 93% to 83% in the dry treatment as the season progressed. Ψ I at midday decreased in both the wet and dry treatments, but the seasonal decline was more pronounced in the latter: seasonal minimum values were -1.19 and - 2.08 MPa, respectively. Plants in the wet treatment maintained turgor (Ψ p ) at about 0.5 MPa throughout the season, whereas values in the dry treatment approached zero towards the end of the season. There was a linear relationship between Ψ p and Ψ I , with Ψ p approaching zero at a Ψ I of -2.0 MPa. Mean daily leaf conductance was consistently higher in the wet treatment (0.46-0.79 cm s -1 ) than in the intermediate and dry treatments (0.13-0.48 cm s -1 ). Conductances in the intermediate and dry treatments were similar, and the lower evapotranspirational water losses in the latter were attributable to its consistently lower leaf area indices (L): L at final harvest was 3.3, 3.3 and 1.9 for the wet, intermediate and dry treatments. Bambara groundnut was apparently able to maintain turgor through a combination of osmotic adjustment, reductions in leaf area index and effective stomatal regulation of water loss.

SUMMARYStands of bambara groundnut (Vigna subterranea (L.) Verde.) were grown in five controlledenvironment glasshouses at the Tropical Crops Research Unit, University of Nottingham, Sutton Bonington Campus, in 1990. Five soil moisture regimes were imposed (one per house), from fully irrigated each week (treatment A), to no irrigation after crop establishment at 35 days after sowing (DAS) (treatment E). Decreasing the amount of water applied resulted in a decline in total dry matter production and harvest index, and a reduction in pod yield from 412 (treatment B) to 0·041 ha-1 (treatment E) at 125 DAS. A maximum leaf area index of 5–4 was achieved by treatments B and C at 90 DAS, resulting in a fractional interception of c. 0·8 of incoming radiation. Total accumulated radiation interception values were 749, 693, 688, 618 and 554 MJ m-2 for treatments A, B, C, D and E, respectively. The efficiency of conversion of the radiation intercepted into dry matter was reduced from 1·41 to 0·50 g MJ-1 by drought.

SUMMARYBambara groundnut (Vigna subterranea (L.) Verde.) is grown throughout southern and western Africa, primarily as a subsistence crop, but agronomic information is scarce. The rates of emergence, flowering and pod production were assessed for 20 weekly sowings between 16 October 1990 and 26 February 1991 at Sebele, Gaborone, Botswana. Daylength during this period changed from 12·7 h to 13·7 h and back to 12·6 h. Mean time to 50% emergence was 111 days while mean time to 50% flowering was 47·5 days. The thermal time to first pod production varied with daylength, decreasing as daylength decreased, to an approximately constant value when daylength after flowering was less than about 12 h. Consequently, the plant size at which pods were produced also varied. The practical implications for date of sowing and plant spacing are discussed.

Bambara groundnut seeds, produced under five levels of irrigation (0–300 mm range) in a suite of controlled-environment glasshouses, were analysed for their proximate composition. There was no significant effect of water treatment on the seed composition. Mean values (g kg–1 DM) of dry matter, protein, starch, free sugars, non-starch polysaccharides and lipids were 8.91, 2.53, 4.28, 0.15, 2.56 and 0.79, respectively. Bambara groundnut protein was deficient in the sulphur amino acids, methionine and cysteine but met the FAO requirements for all other essential and non-essential amino acids. Of the non-starch polysaccharides (NSP) of bambara groundnut, the ratio of insoluble to soluble fractions was 57:43. Cellulose accounted for 36% of the total NSP. Approximately 12.8% of the total NSP was resistant to acid and enzymic hydrolysis. This fraction was attributed to lignified seed coat material and low molecular weight phenolic compounds. Tannin complexes capable of precipitating 142.6 g BSA (bovine serum albumin) per kg seed and trypsin inhibitor activity (13 g kg–1 DM) were measured in all five bambara groundnut treatments. The presence of these anti-nutrients may have implications for the bio-availability of bambara groundnut protein.