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.) Walp) is an important protein-rich grain legume of major economic importance. It is widely grown by small-scale farmers in the arid and semi-arid regions of the world where it is cultivated for its leaves, fresh immature pods and dry grains. However, it is also an underutilized grain legume. In sub-Saharan Africa where most of the cowpea is produced, drought stress is one of the major factors limiting its productivity. Despite the inherent capacity to survive drought stress, several cowpea varieties are affected by mid and late season drought. Therefore, varieties with a higher tolerance to drought stress are required to obtain higher and more stable yields. The objectives of this study were: (i) to determine morphological responses of four dual-purpose cowpea landraces to water deficits during vegetative and reproductive stages (ii) to determine physiological responses of four dual-purpose cowpea landraces to water deficits and recovery during the reproductive stage (iii) to determine yield performance of cowpea landraces after recovery from water stress and how this relates to (ii) above. Four cowpea landraces namely; Lebudu, Lehlodi, Sejwaleng and Morathathane collectedfrom Kgohloane and Ga-Mphela villages, Limpopo Province, South Africa were used in the study. Pot experiments were conducted under glasshouse conditions at the Controlled Environment Facility (CEF), University of KwaZulu-Natal. The first pot experiment evaluated the morphological responses of four cowpea landraces to water stress and recovery. The study was conducted as a single factor experiment laid out in randomized complete block design (RCBD). The treatments (four cowpea landraces) were each planted in 40 pots giving a total of 160 experimental units (drained polyethylene pots with a 5 litre capacity). Each plant in each pot was treated as a replicate. Plants were well-watered until the formation of six fully expanded trifoliates, then irrigation was withheld for 28 days to simulate drought stress during the vegetative growth. The imposition of drought stress was terminated by re-watering all plants after 28 days. The cowpea plants were re-watered sufficiently and allowed to grow until the four landraces reached 50% flowering stage. Watering was withheld again at 50% flowering for a two-week period for all the four landraces to simulate drought stress during the reproductive growth. The second experiment was conducted to investigate physiological responses of the four cowpea landraces to water stress during the reproductive stage. The experiment was laid out as a 4 x 2 factorial treatment structure in randomized complete design (CRD) with the following three factors: cowpea landraces – 4 levels (Lebudu, Lehlodi, Sejwaleng and Morathathane), water regimes – 2 levels (stressed and well-watered) treatment combinations each replicated 20 times (20 pots each containing one plant) giving a total of 160 experimental units (drained polyethylene pots with a 5 litre capacity). Data on morphological responses were collected and included: number of green vs. senesced leaves, visual assessment of leaf greenness, stem, branch greenness and survival percentage. Physiological responses to water stress were determined during the reproductive stage and included: leaf water potential, relative water content, stomatal conductance, proline content, chlorophyll content, carotenoid content, chlorophyll a content, phenolics (free and membrane-bound), total antioxidant capacity and chlorophyll fluorescence parameters (Fv/Fm). Genstat 14th edition (VSN International, UK) was used to perform analyses of variance (ANOVA) and differences between means were determined by the Least Significant Differences (LSD) at the 5% probability level. Landraces showed different morphological responses during both vegetative and reproductive growth stages. Lebudu, Lehlodi and Sejwaleng displayed a strong ability to maintain stem greenness longer as compared to Morathathane during vegetative growth. Lebudu delayed leaf senescence more than other landraces; no differences in survival were observed. All landraces survived for 28 days without water and resumed growth after re-watering. During the reproductive stage, Lebudu displayed a strong ability to maintain leaf, branches and stem greenness longer and showed relatively higher tolerance to drought stress compared to other three landraces. Water stress caused a decline in leaf water potential, relative water content, carotenoid content, chlorophyll content, stomatal conductance and increased proline content, phenolics, chlorophyll a content, total antioxidant capacity and while chlorophyll fluorescence parameter, Fv/Fm, was not affected. All landraces maintained higher relative water content above a critical threshold with Sejwaleng maintaining a significantly higher RWC (69%) than Lehlodi, Lebudu and Morathathane. Morathathane developed a more negative leaf water potential at maximum stress than Lebudu, Lehlodi and Sejwaleng. Stomatal closure was observed in all cowpea landraces during water stress, but re-opened after re-watering. Chlorophyll content was considerably reduced in Morathathane as compared to Lebudu, Lehlodi and Sejwaleng. No significant differences were observed between the cowpea landraces with respect to carotenoid content at maximum stress. Chlorophyll a content increased significantly for Morathathane as compared to Lebudu, Lehlodi and Sejwaleng. High accumulation of proline was observed for Lebudu, Lehlodi and Morathathane as compared to Sejwaleng, which showed a very slow accumulation of proline. Lebudu, Lehlodi and Sejwaleng showed an increase in phenolic compounds while a decline was observed for Morathathane. Total antioxidant capacity (TAOC) was high in all cowpea landraces during water stress. Also, all chlorophyll fluorescence parameters showed that cowpea landraces had efficient photo-protection mechanisms during drought stress. After re-watering, relative water content, leaf water potential, stomatal conductance, chlorophyll content, carotenoids, chlorophyll a, proline content and TAOC recovered and reached the same level as that of well-watered plants. All four landraces were re-watered after the imposition of stress and above ground biomass, pod mass and number and seed yield determined. Although there was a reduction in the total above-ground biomass, pod mass and number in all four landraces under water stress compared to the well–watered treatment; this was not statistically significant (P > 0.05). Furthermore, no significant differences (P > 0.05) were observed between the four landraces with respect to seed yield under stressed and well-watered conditions. This study established that cowpea landraces vary with respect to the various morphological and physiological adaptive mechanisms in response to water deficits. Such adaptive mechanisms probably ensure their survival under severe water stress conditions until the next rainfall and therefore allowing them to produce reasonably relatively higher leaf and seed yield. Detailed knowledge of these mechanisms in the landraces could be useful in the genetic enhancement and breeding for drought tolerance in the existing cowpea germplasm. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2013.

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.