Cowpea (Vigna unguiculata L.) is an important legume, especially in the hot, dry tropics and subtropics of sub-Saharan Africa. It has been widely reported to be drought tolerant. Cowpea is a highly nutritious, multi-purpose crop, used as a leafy vegetable and grain legume with potential to contribute to food security in marginal areas. However, the crop is still classified as a neglected underutilised species; legume research focus has been mainly devoted to established legumes such as common bean and soybeans. There is a need to collect empirical information on cowpea which could be used to advise farmers on management strategies. This study evaluated cowpea responses to water stress under controlled and field conditions. Initially, two cowpea varieties (Brown and White birch) were evaluated for seed quality using the standard germination that was laid out in a completely randomised design and each variety was replicated for times. Electrolyte conductivity test was also performed under laboratory conditions. Thereafter, a pot trial was conducted to evaluate cowpea response to water stress imposed at different growth stages under varying growth temperatures. The pot trial comprised three factors: temperature [High (33/27ºC), Optimum (27/21ºC) and Low (21/15ºC)], water regimes (no stress, terminal stress, intermittent stress – vegetative and intermittent stress - flowering) and cowpea varieties. Lastly, a field trial was conducted to evaluate cowpea production as well as the effect of sequential leaf harvesting on yield under irrigated and rainfed conditions. The field trial was laid out as a split-plot design, with water regime (irrigation vs. rainfed) as main factors, cowpea varieties as sub-factor and sequential harvesting (no harvest, harvested once and harvested twice), replicated three times. All treatments were arranged in a randomised complete block design. Results of the initial study showed that germination capacity and vigour of cowpea varieties were significantly different (P < 0.001). White birch had higher electrolyte leakage than Brown birch. Pot trial results showed that cowpea growth (leaf area, leaf number and plant height) was vigorous in the high temperature regime compared with optimum and low temperature regimes. Chlorophyll content index was higher under high temperature relative to optimum and low temperature regimes, respectively. Under low and optimum temperature regimes, cowpea growth was stunted; cowpea failed to flower and form yield. Whereas, under high temperature regime, cowpea growth was vigorous hence flowered and formed yield. Vegetative growth was more sensitive to water stress than flowering stage. Terminal stress and stress imposed during flowering resulted in increased proline accumulation relative to no stress and stress imposed during vegetative growth. Harvest index was lower when water stress was imposed during vegetative relative to flowering stage. Field trial results showed that cowpea growth was sensitive to water stress. Plant height, leaf number, chlorophyll content index and stomatal conductance were lower under rainfed relative to irrigated conditions. Sequential harvesting of leaves had no significant effect on cowpea yield. It is concluded that tropical temperature conditions are most suitable for cowpea production; the controlled environment study showed best crop performance under 33/27ºC. In the context of varieties used for the present study, vegetative growth was the most sensitive stage to water stress. Cowpea performed better under rainfed relative to irrigated conditions with respect to yield formation. Low temperature was found to be more limiting to cowpea growth, development and productivity compared with water stress. Whereas, under high temperature conditions, water stress was more limiting to plant growth and productivity. White birch may be used as a dual purpose crop due to its ability to produce reasonable grain yield regardless of defoliation. / Thesis (M.Sc.Agric)-University of KwaZulu-Natal, Pietermaritzburg, 2012.

Cowpea (Vigna unguiculata. L) is an important African crop. However, it is also an underutilized grain legume. Consequently, there is not enough research data on cowpea seed physiology. Whereas there is evidence of cowpea being a drought tolerant crop, there is no evidence to associate plant drought tolerance with seed quality in response to water stress. This study sought to understand the effect of production site and water stress on cowpea seed quality development with respect to germination capacity and vigour. Patterns of raffinose family of oligosaccharides (RFO) during seed development to mature dry stage were used to physiologically relate seed performance to water stress. The effect of water stress and exogenous ABA on the accumulation of stress LEA proteins (dehydrins) in relation to seed quality development and germination was investigated. RFOs are known for their roles in desiccation sensitivity but no studies have shown their significance in cowpeas. Seeds of six cowpea cultivars were produced at two distinct growth sites characterised by irrigated and dry land conditions. The seeds were assessed during six developmental stages, for water content, dry matter accumulation, and performance. Harvested seeds were then planted in a pot experiment under controlled conditions to examine the effect of water stress on seed quality development and data collected during three developmental stages. Harvested seeds from the pot experiment were subsequently analyzed for changes in RFO accumulation during development using gas chromatography. The seeds were also used to investigate the effect of water stress and ABA on the accumulation of stress LEA proteins (dehydrins) in relation to seed quality development in cowpea. In addition, this study evaluated the use of image analysis as a method that can be used to objectively determine seed coat colour variation in cowpea. Statistical variation in individual seed’s solute leakage for cowpea cultivars differing in seed coat colour and produced under different environmental conditions was explored and correlations were done between seed conductivity test with other aspects of seed performance during germination. Furthermore the results of the conductivity test were compared with accelerated aging test, in relation to seed performance. The study provided evidence that cowpea seed lots produced under different environmental, and possibly management conditions may not differ with respect to seed quality as determined by germination capacity and vigour. However, significant differences between sites with respect to seed maturation patterns determined by water content and dry matter accumulation were observed. Adverse maternal environmental effects on the subsequent performance of seeds in a drought tolerant crop may not necessarily lead to poor performance. Cultivar differences in response to simulated drought conditions at the whole plant and tissue level can be considerable and highly variable; however, these differences may not have adverse effects on the germination and vigour of the seeds. Drought avoidance mechanisms at the whole plant level in cowpea are quite efficient in allowing the species to adapt to simulated drought conditions. These mechanisms may allow the cowpea cultivars to maintain metabolism and restore conditions for their continued growth under water stress; and produce few seeds of high germination capacity and vigour. Stachyose was found to be the predominant member of the raffinose family of oligosaccharides in cowpea. It is suggested that stachyose accumulation could be used as an indicator of stress tolerance in cowpea. However, the relationship between RFO concentration and the acquisition of desiccation remained as a matter of speculation in the present study and is still generally inconclusive. There was no evidence to suggest the acquisition of maximum desiccation tolerance is associated with maximum seed vigour. It is suggested in cowpea, which is drought tolerant, that maximum vigour does not necessarily imply the acquisition of maximum desiccation tolerance; rather there is a minimum level of desiccation tolerance that is required for the development of optimal seed vigour. The use of an in vivo approach in the study of LEA function in cowpea enabled the accurate comparison of two different groups of LEA proteins in developing cowpea seeds under conditions of water stress and in relation to germination and vigour. Both group 1 LEA and group 2 LEA (dehydrin) were shown to increase in concentration in response to water stress. In addition group 1 LEA protein was observed to be relatively abundant in cowpea seeds. A maternal influence on LEA protein gene expression under conditions of water stress, which may induce dehydrin accumulation vii during the earlier stages of seed development, was implied by the observation that dehydrin-like proteins were induced after two weeks of development in cowpea plants subjected to stress during the vegetative phase. In addition, the exogenous application of ABA delayed radicle protrusion; this was associated with a delay in the disappearance of LEA proteins and is suggestive of a relationship between LEA protein accumulation and the acquisition of desiccation tolerance. The study has demonstrated that image analysis can objectively discriminate seed coat colour variation in cowpea. Dark coloured seeds in general performed better than light coloured seeds; however seed coat colour was not always associated with better performance. A newly developed Aging Stress Differential Index (ASDI) has been used in this study to demonstrate a link between seed coat colour and sensitivity to water stress. The ASDI correlated well with the observations relating stress tolerance to stachyose accumulation. The skewed distribution patterns in individual electrical conductivity and the presence of extreme values may have implications with respect to the suitability of using standard statistical analyses which compare mean values to evaluate such data. In addition variation in individual electrical conductivity may also be influenced by cultivar differences and the chemical composition of the seed coat. Therefore associations between seed coat colour and electrical conductivity as a measure of performance should be treated with caution. The AA test does reflect changes in seed vigour, however ranked electrical conductivity values after AA did not consistently reflect differences in seed performance between cultivars and sites, and they did not correlate well with other aspects of performance. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2007.

Cowpea yields in Mozambique can be increased through breeding farmers’ accepted cultivars with drought tolerance and stability across environments. A study was conducted in the southern region of Mozambique to: (1) determine farmers perceptions on major constraints limiting cowpea production and identify preferences regarding cultivars and traits, (2) determine the variability of selected cowpea germplasm for drought tolerance, (3) determine the gene action controlling drought tolerance, yield and yield components in cowpea, and (4) assess the genotype × environment interaction and yield stability of cowpea genotypes under drought-stressed and non-stressed conditions. The study on farmers’ perceptions about the major constraints limiting cowpea production and preferences regarding cowpea cultivars and traits established that cowpea was an important crop, cultivated for its grain, leaves and fresh pods for household consumption and the market. The study revealed that cowpea grain and leaves were equally important across the three districts in the study. Differences in accessibility to markets between districts influenced the ranking of grain and leaves among districts. Grain was more important in Bilene and Chibuto districts which are situated far from the major urban centre, Maputo, while leaves were more important in Boane district which is near the major market of Maputo. Fresh pods were important in Bilene district which is situated along the major highway connecting Maputo and other provinces. Drought was the most important production constraint followed by aphids, bruchids and viral diseases. The criteria used by farmers to select cowpea varieties included high grain and leaf yield, large seed size, earliness, smoothness of the testa and potential marketability of the variety. The implication of this study is that different types of varieties need to be developed for different areas. Dual-purpose or grain-type varieties need to be developed for areas situated far away from the major markets while varieties for leaf production need to be bred for areas near major markets. During the breeding process, a selection index needs to be adopted whereby drought tolerance, high grain and leaf yield, large seed size, smooth testa, earliness, aphids and bruchids resistance should be integrated as components of the index. High grain yield should receive high weight for varieties developed for areas located far from major markets while high leaf yield would receive high weight for varieties developed for areas located near major markets. The study on variability of cowpea germplasm collections for drought tolerance revealed wide genotypic variability among the tested germplasm. Biplot displays indicated that the genotypes could be grouped into four categories according to their drought tolerance and yielding ability as indicated below: high yielding-drought tolerant (group A), high yielding-drought susceptible (group B), low yielding-drought tolerant (group C), and low yielding-drought susceptible (group D). Examples of high yielding-drought tolerant genotypes were Sh-50, UC-524B, INIA-24, INIA-120, IT96D-610 and Tete-2. Stress tolerance index was the best criterion for assessing genotypes for variability in drought tolerance because it enabled the identification of high yielding and drought tolerant genotypes (group A). The assessment on gene action controlling drought tolerance (stay-green), yield and components indicated that both additive and non-additive effects were involved in controlling all of these traits. Additive gene action was more important than non-additive gene affects in controlling stay-green, days to flowering, number of pods per plant, number of seeds per pod and hundred seed weight. Under no-stress conditions, additive gene action was more important than non-additive gene action while under drought-stressed conditions, non-additive gene effects were more important than additive gene effects. Stay-green can easily be assessed visually in early segregating populations while yield and yield related traits cannot. Hence, selection for drought tolerance using the stay-green trait would be effective in early segregating generations while selection for yield and number of pods per plant would be effective in late segregating generations. Selection for yield could be conducted directly under no-stress conditions and indirectly using the number of pods per plant under drought stress conditions. Genotype INIA-41 would be the most desirable to use as a parent for drought tolerance and IT93K-503-1 would be the most desirable to use as a parent for drought tolerance and yield. The assessment on genotype × environment interaction and cowpea grain yield stability for forty-eight (48) cowpea genotypes grown under drought-stressed and non-stressed conditions indicated that cross-over genotype × environment interactions were present for yield indicating that genotypes responded differently to varying environmental conditions. Genotypes adapted to specific environmental conditions could be identified. Genotypes IT-18, INIA-51, INIA-51A and Nhavanca were adapted to non-stressed environments that were either drought stressed or non-stressed while VAR-11D was adapted to low yielding, stressful environments. Genotypes INIA-23A, INIA-81D, INIA-24, INIA-25, INIA-16 and INIA-76 were high yielding and stable while genotypes IT-18, INIA-51, INIA-51A, Nhavanca and VAR-11D were high yielding and unstable. Genotypes Bambey-21, INIA-36, INIA-12 and Monteiro were consistently low yielding and stable except INIA-12 that was consistently unstable. Chókwè was a high yielding environment and suitable for identifying high yielding genotypes but not ideal for selection because it was not representative of an average environment while Umbeluzi was low yielding and not ideal for selection. Overall, the study revealed that genetic improvement of drought tolerance and yield would be feasible. Potential parents for genetic improvement for yield and drought tolerance were identified. However, further studies for assessing yield stability of cowpea genotypes are necessary and could be achieved by including more seasons and sites to get a better understanding of the genotype × environment interaction and yield stability of cowpea in Mozambique. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2010.