1. Biotic Stresses
1.1. Bacterial Diseases
Bacterial disease is the most economically important biotic stress in cowpea production in SSA countries. Among them, cowpea bacterial blight (CoBB) and bacterial pustules are some of the most severe bacterial infections of cowpea, causing severe damage
[1][2]. CoBB symptoms start with small water-soaked spots on leaves, which enlarge to irregular brown necrotic lesions surrounded by yellow haloes, leading to premature leaf drop
[1]. The pathogen also invades the cowpea stem, causing canker symptoms in susceptible plants
[2]. It generally infects all growth stages of the cowpea plant: seedling, vegetative, flowering, and podding stages, and all plant parts, including leaves, pods, and seeds
[3]. Frequent evaluation of cowpea genetic materials and resistant line development is the best option for achieving desirable resistance gene sources for use in bacterial resistance through breeding programs.
1.2. Viral Diseases
Globally, cowpea is infected by more than 140 viruses, but only nine were reported as economically important
[4]. Cowpea aphid-borne mosaic virus (CABMV), cowpea golden mosaic virus (CPGMV), Southern bean mosaic virus (SBMV), Sunhemp mosaic virus (SHMV), blackeye mosaic virus (BICMV), cucumber mosaic virus (CMV), cowpea mottle virus (CMV), cowpea yellow mosaic virus (CPMV), and cowpea mild mottle virus (CPMMV) are the most common seed-borne viral diseases
[5]. Viral diseases can cause a yield loss of 10% to 100%, depending on the time and the severity of infection in cowpea
[6]. The aphids suck the cowpea sap, which might affect the physiological processes and mineral element transportation in the plant system and consequently affect its concentration
[7]. It is also a vector in the transmission of viruses
[8]. For tackle the existing viral disease problem, intensive evaluation of cowpea genotypes with multiple viral infections under greenhouse and hotspot production areas are the best option to develop resistant varieties. In addition, using biotechnology that will result in the production and deregulation of virus-resistant cowpeas through coat-protein gene transfers should be intensified
[4].
1.3. Root-Knot Nematode
Root-knot nematode,
Meloidogyne incognita, is a severe pest and a major constraint on cowpea production in most growing areas of the world, causing 80 to 100% yield losses
[8][9][10][11][12][13][14][15][16][17][18][19][20][21][22][23][24].
M. incognita and
M. javanica are the major species found on cowpea in most growing regions
[9]. Damage symptoms of root-knot nematodes include patches of stunted and yellowed plants. Severe damage can lead to reduced numbers of leaves and buds
[25].
1.4. Parasitic Weeds
Parasitic weeds,
Striga gesnerioides (Willd) Vatke ex Engl and
Alectra vogelii (Bent), are serious threats to cowpea production in Africa
[6][8]. Weeds reduce cowpea yield and quality by competing for light, space, water, soil nutrients, and carbon dioxide
[9][10]. In total, yield losses caused by parasitic weed infestation alone in cowpea ranges from 73 to 100%
[10][11][23]. These weeds may also reduce productivity by releasing allelopathic compounds into the environment
[16] and by providing a conducive environment and serving as a vector for insect pests and viruses
[17].
Completely removing these parasitic weeds in cowpea production is too difficult because the seeds can remain viable in the soil for up to 20 years
[6][15]. Therefore, the best and most sustainable option to address this critical problem is to breed cowpea varieties which are resistant to these weeds, using multiple smart and agro-biotechnology techniques that could be deployed sustainably.
1.5. Insect Pests
Cowpea is attacked and damaged by insect pests in all stages of growth
[19]. Insects are the most challenging threat to cowpea production and productivity because they occur at pre-flowering, post-flowering, and storage
[20]. Seed com, maggot, cutworm, aphids, and leafhopper occur at the pre-flowering (seedling) stage. Aphids, leaf miners, and thrips are also active insect pests at the flowering and post-flowering stage, and aphids, bean fly, bean pod borer, leaf miner, and thrips are the common insect pests at the reproductive stage (grain filling period) of cowpea
[21]. Among the field pests of cowpea, aphid [
Aphis craccivora (Koch)] is an important vegetative stage pest of cowpea in Africa but also occurs at other growth stages. Both nymphs and adults suck plant sap and cause serious damage from the seedling to the pod bearing stage
[22]. Aphids cause damage through secretion of honeydew, which promotes the growth of sooty molds and other fungi on leaves, curling of leaves and delayed flowering, shriveling of pods and, as a result, reduced photosynthetic processes and rates, finally resulting in overall yield reduction
[23]. It affects the crop by directly sucking its sap. Their feeding on cowpea causes cupping of the leaves, crinkling, defoliation, and stunted growth
[24]. Another serious effect of aphids is the ability to transmit the aphid-borne mosaic virus. Affected plants show a green vein banding of the leaves
[26]. Molecular and phenotypic screening of cowpea
[11] identified cowpea genotypes with good resistance to aphids, which can be used as a source of resistance genes in breeding new aphid-resistant cultivars.
2. Abiotic Stress
2.1. Drought
Cowpea is known to be drought-tolerant compared to other crop plants
[27][28]. Among the abiotic factors, drought has been identified as a significant limitation, restricting cowpea production and productivity
[27]. In the arid and semi-arid tropics, the productivity of cowpea could be hampered by erratic rainfall at the beginning and towards the end of the rainy season
[29]. Drought leads to adverse influences on cowpea growth, development, and reproduction ability
[30], limiting the crop’s yield and productivity
[31]. This crop plant is a robust legume; nevertheless, drought always affects yield, especially during the reproductive and seed-filling period
[32]. This causes a substantial reduction in grain yield and biomass production
[28]. The influence of drought varies and depends on the intensity, developmental stage, and duration of stress and the adaptive strategy that the plant possesses to tolerate this stress
[28][33]. Cowpea suffers from drought stress due to erratic rainfall due to climate change, resulting in a yield loss of up to 35–69%
[34]. To provide a better solution to drought stress in cowpea production in SSA, a comprehensive evaluation and characterization of cowpea genotypes for developing drought-tolerant varieties using physiological, biochemical, and molecular approaches could be exploited. Understanding drought tolerance gene expression will further advance tolerance breeding.
2.2. Salinity
Salt stress is one of the most significant abiotic constraints to cowpea crop productivity and severely hampers crop production, especially in arid and semi-arid areas. Cowpea is unfavorably affected by salinity stress at seed germination and seedling stages, and growth and vigor are reduced, which is exacerbated by climate change effects
[35]. Salinity stress ultimately reduces the yield (leaves, immature pods, and grain weight and quality) of cowpea, while other vegetative growth traits are also adversely affected
[36][37]. In addition, salinity reduces the ability of cowpea crops to take up water and soil-plant nutrients, leading to growth reduction and metabolic changes similar to those caused by low soil moisture stress
[38]. Furthermore, it reduces lipid peroxidation and leads to destructive oxidation, which in turn causes damage to the key plant biomolecules
[39]. Salt stress is a complex trait, and it is associated with other agronomic and biochemical traits of cowpea. Therefore, for developing salt stress-tolerant cowpea genotypes, integrative (biochemical, molecular, and conventional) breeding approaches are the best solution for tackling the existing problems in SSA.
2.3. Heat Stress
Heat stress is a crucial abiotic stress that significantly affects the growth and yield of cowpea
[40]. The effects of heat stress on yield and yield-contributing traits include flower and leaf drop, poor pollen fertility and germination, low pod setting, low plant biomass, low harvest index, poor pod filling, and low seed weight and yield
[41][42]. Furthermore, physiological and biochemical traits of cowpea negatively affect the photosynthetic apparatus, such as impaired photo-assimilation, inhibited N2 fixation, increased leaf senescence, decreased canopy temperature, and leaf relative water content
[42][43][44]. When the night temperature reaches about 16 °C, cowpea flowers abort due to poor pollen development, which causes a 4 to 14% loss in leaves, immature pods and grain yield and quality
[45]. In general, using conventional, physiological, phenomic, functional genomic, proteomic, and metabolomic breeding techniques
[44][46][47][48][49] and viable approaches to developing heat stress tolerance and to sustain cowpea yield under increasing high-temperature stress, the cowpea germplasm needs to be screened to identify ‘stress adaptive’ traits across various gene pools.
2.4. Low Soil Fertility
Phosphorus is essential for cowpea production in many tropical African soils with the inherent low soil availability of phosphorus
[50]. Cowpea does not require too much nitrogen fertilizer because it fixes its nitrogen from the air using the nodules in its roots
[51]. However, phosphorus is critical to cowpea yield because it stimulates growth, initiates nodule formation, and influences the efficiency of rhizobium-legume symbiosis
[52]. Therefore, cowpea requires more phosphorus than nitrogen in the form of single super phosphate
[53]. In addition, it is required in large quantities in young cells such as shoot and root tips where metabolism is high and cell division is rapid
[54]. It also aids in flower initiation and seed and fruit development
[55].