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Cowpea cultivation constraints and breeding in Europe: Comparison
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Cowpea (Vigna unguiculata (L.) Walp.) is a legume with a constant rate of cultivation in Southern European countries. Consumer demand for cowpea worldwide is rising due to its nutritional content, while Europe is attempting to reduce the deficit in the production of pulses and invest in new, healthy food market products. Cultivation and utilization of cowpea plant genetic resources (PGRs), including landraces, in breeding programmes with the implementation of classical and modern breeding techniques could promote sustainability of cropping systems and alleviate the negative effects of climate change.

  • abiotic stresses
  • biotic stresses
  • breeding methods
  • crop wild relatives
  • landraces

1. Introduction

Cowpea (Vigna unguiculata (L.) Walp.) (2n = 2x = 22) is an important legume species, both for its consumption as food and as animal feed worldwide, especially in semi-arid tropical and desert regions [1],[2]. It is an excellent source of vitamins, antioxidants, fiber, trace elements and other nutrients [2],[3] and plays an important role in malnutrition avoidance in the least developed countries (LDCs) where it is mainly cultivated [4],[5]. Almost all its above-ground plant parts are consumed [6]. In addition to its mature dry seeds, its leaves, green pods and green seeds are consumed in various countries [5],[7],[8],[9],[10]. It is also used for flour [3],[11],[12],[13], as its seeds contain a high protein content (23–32%) compared to many other legume species [14],[15].

2. Botanical taxonomy, Origin and Spread

The genus Vigna belongs to the legume family and currently includes approximately two hundred species [16], among them ten cultivated species, such as cowpea (Vigna unguiculata (L.) Walp.) and green mung bean (Vigna radiata (L.) Wilczek) [17]. The species of the genus are grouped into six subgenera (Vigna, Ceratotropis, Plectotropis, Sigmoidotropis, Lasiosporon and Haydonia) [18],[19], while more recent studies through molecular phylogenetic analyses recommended the removal of the subgenus Sigmoidotropis [20],[21]. Cowpea is classified in the Catiang section of the subgenus Vigna [22]. The cultivated types of the species (Vigna unguiculata ssp. unguiculata var. unguiculata) are further categorized into four cultivated groups (cultivar groups, cv.-gr.): sesquipedalis, textilis, biflora and unguiculata [23],[24], while Pasquet [25] proposed the introduction of one more cultivated group, melanophthalmus. In many cases, these cultivar groups are considered by some researchers as separate subspecies and not as cultivar groups of a specific subspecies [26].

Cowpea (Vigna unguiculata (L.) Walp.) was domesticated in Africa before 1500 B.C. [27]. Southeast Africa or West and Central Africa [28],[29],[30],[31],[32] are suggested as the primary center of the species origin. The existence of a parallel double center of cowpea domestication was mentioned lately [33]. It seems that cowpea was first introduced into Asia during the Neolithic period, around 1500 B.C. [34],[35], and in India, a secondary center of origin for the species was formed. Upon entering Asia, cowpea encountered different climatic conditions, and after selection for fresh pod consumption, the cultivated group V. unguiculata ssp. unguiculata cv.-gr. sesquipedalis was formed [33],[36]. This cultivated group was later introduced into the European continent. Along with the spread of cultivar group V. unguiculata ssp. unguiculata cv.-gr. sesquipedalis in the European area, the spread of the African cultivated Vigna (Vigna unguiculata ssp. unguiculata cv.-gr. unguiculata) probably also took place [37]. As testified by the texts of Theophrastus, cultivation of the “bean” was familiar to the ancient Greeks in 300 B.C. [38]. The two cultivated groups later spread from Europe to South America during the 17th century A.D.[37], and then to the USA during the 18th century A.D. [39].

Today, cowpea is cultivated in areas of Southern Europe such as Greece, Italy, Spain, Cyprus, Croatia, Portugal and Serbia, Bosnia and Herzegovina, and North Macedonia [40],[41],[42],[43],[44],[45], but is less widespread in Central Europe [38]. Cultivated areas for cowpea dry beans production are also reported in Slovenia and Hungary [44]. In total, cowpea corresponds to the 0.3% of pulses production in Europe and the 0.3% of world cowpea production for dry seed, reaching up to 23,825.22 ton in 2021 [44],[46]. However, not much data are available regarding cowpea production for fresh pod production despite that its production was enhanced in Southern European countries lately [10],[47]. 

3. Constraints and breeding methods

Cowpea faces a multitude of biotic and abiotic factors that negatively affect its productivity worldwide [48]. Regarding breeding classical methods are used mainly such as pedigree selection, mixed population selection and single seed-derived selection and pure line selection [5],[49],[50],[51]. Backcrosses are usually applied To introduce a desired trait into an already adapted parent [52],[53],[54]. Modern and assisted techniques are also implemented, like induced mutagenesis, tissue culture, genetic modification, introgression breeding, use of molecular markers, quantitative trait loci (QTLs) identification and genome-wide association studies (GWAS) [36],[55],[56],[57],[58],[59],[60],[61],[62],[63],[64],[65],[66],[67],[68],[69].

In Europe cowpea cultivation area is increasing [10] as it is considered a drought and high temperatures tolerant plant species in comparison to other legumes. Recently, cowpea fresh pods and green seeds started to be investigated as new products for the market [10], while more intense efforts are made to increase pulse production and grain legume-based products in Europe [70]. Cowpea cultivation in temperate climates lasts from late Spring to early Autumn. Although the area is characterized by mild climatic conditions and the crop is not subjected to such adverse conditions as in Africa [5], cowpea confronts with a plethora of stresses and yield limiting factors regarding seed and fresh pod production. Exploitation of nutrient content of its fresh pods and seeds is also important worldwide as many substances are essential for humans [71]. Identification of genetic material available and suitable regarding adaptation and tolerance to the main cowpea restrictive factors is the first step for breeding due to the narrow genetic base that characterizes breeding varieties [72]. Wild relatives, exotic germplasm and landraces are sources of hidden diversity. Landraces form the main variability source for cowpea, as there are incompatibility difficulties and production of non-fertile hybrids while inter-crossing with crop wild relatives [45].

Cowpea confronts with various abiotic stresses i.e., high temperatures [73], drought especially during flowering and pod setting periods [74],[75], photoperiodic requirements [48],[76] and soil limiting factors [77] such as salinity and sodicity problems [78]. In Southern Europe, with reduced water availability looming worldwide [14], searching for drought-tolerant genetic material along with production stability under limiting water conditions is of primary importance. Numerous genotypes and accessions, including landraces, have been characterized and evaluated with the goal of being a genetic resource for drought [74],[79], heat and salinity tolerance [80],[81],[82],[83],[84],[85],[86],[87]. No extensive screening and evaluation of cowpea landrace material have been done with regard to other abiotic stresses. However, several cowpea genotypes and cultivars have been identified [82],[83],[88],[89], and breeding lines with high temperature tolerance have been developed [86],[90]. Landraces and other cultivated material were screened and appeared also to be promising in cultivation under calcareous and high alkaline soils [42],[91].

Among the screened genotypes for biotic stress factors some landraces and wild cowpea accessions presented resistance to aphids, like twelve Greek cowpea landraces to Aphis craccivora was highlighted as they found to possess the allele CP-171-172 indicative for aphid resistance of the TVu-2876 genotype [45]. Two landraces were also identified by Nyarko et al. [92] to present weevil resistance. Nematodes resistance was also observed in twelve wild and landrace accessions by Dareus et al. [93], while many resistant landraces along with genotypes and varieties to viruses have also been identified [94],[95].

European cowpea landraces have been studied, revealing interesting material regarding their yield potential. In a collection of 48 cowpea accessions evaluated by Stoilova and Pereira [96] including landraces, a landrace from Bulgaria named “A4E007” was among the genotypes selected to be included in a breeding programme aiming to increase seed yield. Martos-Fuentes et al. [76], evaluating, at three locations and two consecutive years, twelve cowpea genotypes, consisting mainly of landraces material from Greece, Portugal and Spain, defined cowpea landraces with high productivity levels such as “BGE038474” in the Cartagena region and “Vg73” at the Elvas site. Cowpea landraces with a European origin have been also evaluated regarding their seed, fresh seeds and green pods proximate and nutrient composition, revealing some promising genotypes such as “Cp5647” and “Cp4877” originating from Portugal that exhibited high total soluble solids (7.6 and 6.5 °Brix, respectively) and titratable acidity of fresh pods [47], “BGE038477” and “BGE038478” from Spain that presented high seed antioxidant capacity (17.78 mg GA g-1 dw and 18.26 mg GA g-1 dw, respectively) and phenolic content [10], “AUA2” from Greece that presented high Ca (6.10 g kg-1), Mg (3.40 g kg-1), S (1.21 g kg-1), B (20.60 mg kg-1) and Zn (64.10 mg kg-1) content of fresh pods and P (5.40 g kg-1) and Fe (70.90 mg kg-1) content of immature seeds [10] as well as “BGE038477”, “BGE038478” and “VG20” that presented high seed protein content, 29.52%, 29.46% and 28.37%, respectively [[10],[42]].

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