Effect of Water Stress on Morpho-physiological Trait of Some Accessions of Bambara Groundnut [Vigna subterranea (L). Verdcourt]

A
Abib Chimene Fanta1
W
Wassouo Alain Felix2
N
Nassourou Maina Antoine1
D
Difo Voukang Harouna1
T
Tchapda Dorothy Tchatchoua2
1Department of Biological Sciences, Faculty of Sciences, University of Maroua, Po. Box. 814 Maroua, Cameroon.
2Department of Agriculture, Animal Husbandry and Derived Products, National Advanced School of Engineering, University of Maroua. Po. Box. 46 Maroua, Cameroon.
  • Submitted21-05-2025|

  • Accepted29-07-2025|

  • First Online 15-09-2025|

  • doi 10.18805/LRF-878

Background: The cultivation of Bambara groundnut in arid zones, such as the Sahelian regions of Cameroon, has received scant attention despite its nutritional and agronomic potential. The Sahelian regions of Cameroon are classified as part of the arid zones of the world, characterized by a rainy season of approximately three to four months in duration. Nevertheless, the cultivation period of Bambara groundnut is approximately four to six months, a duration that hinders its cultivation. The objective of this study is to assess the impact of water stress on the morpho-physiological traits of select Bambara groundnut accessions cultivated in the Sahelian regions of Cameroon.

Methods: The experiment was executed in accordance with the fisher bloc design, employing four replicates. The water stress was induced using distilled water in pot experiments to monitor thirteen (13) morphological traits and six (06) physiological traits.

Result: The results demonstrated significant variability among the accessions under water stress. Accession V2 exhibited the greatest resistance, characterized by diminished leaf areas and reduced stomatal number. The findings demonstrated a consistent increase in relative water content, accompanied by a decrease in water loss and transpiration. This observation suggests a reduced response to stress. The genetic material of the accession exhibited a range of morphological responses. The considerable impact of water stress was evident on the 100-pod weight produced per accession. Accession V1 exhibited a significant response to water stress, resulting in the production of the smallest pods (22 g). Accession V2 demonstrated the least susceptibility to the 100-pod weight (26 g), exhibiting the highest weight among the other accessions. The evaluation of the 100 seeds’ weight revealed that accession V2 exhibited a significantly higher seed weight compared to the other accessions. The grain yield in the accession V2 was higher for both sites (65.39±1.15 and 65.72±0.50). Principal component analysis demonstrated a positive correlation between stress and the weights of 100 pods and seeds (r = 0.956). The results obtained underscore the capacity of accession V2 to adapt to the environmental conditions prevalent in the Sahelian regions of Cameroon.

Global population growth over the past few decades is generating enormous needs, the most notable of which in sub-Saharan Africa are energy and food demand. in fact, the report published by the FAO, shows the absolute number of people suffering from hunger and under nourishment reached almost 821 million in 2017, compared with around 804 million in 2016, (Salomon et al., 2023), Indeed, in developing countries, 12.9% of the population is undernourished and malnutrition is the cause of 45% of deaths in children under 5.
       
[Vigna subterranea (L.) Verct], commonly known as Bambara groundnut, is a plant indigenous to Africa and is one of the often-neglected subsistence crops in Africa (Soumare et al., 2022). It is a minor food legume adapted to a variety of stressful climatic and ecological conditions (Onwubiko et al., 2011). It is a calorie-rich plant food, with an energy value of around 387 Kcal/100 g. It is also rich in vitamins, minerals, proteins (20-25%), lipids (6-8%) and carbohydrates (50-57%) (Gbaguidi et al., 2015). Bambara peanut seeds are used as human food as well as for the treatment of certain pathogens (Gbaguidi et al., 2015). However, many plant species grown around the world for food production are considered minor crops compared to other food crops considered major food crops (rice, maize, wheat, sorghum, yam, plantain, cassava, etc.) which make up almost the entire diet in Africa (Bricas et al., 2009). However, these minor crops play a crucial role in food security, nutrition and generate income for many rural populations (Ouoba et al., 2018). Despite its nutritional qualities, Bambara groundnut is on the list of neglected species in varietal breeding programs (Tadele Zerihun, 2009). However, the plant offers enormous potential for improvement, thanks to its genetic diversity found in thousands of accessions harvested throughout the world. (Djè et al., 2005); (Ndiang et al., 2012). In Cameroon for instance, very little information is currently available on the genetic and morpho-physiological diversity of this species.
               
In the arid regions of Cameroon, water scarcity and precarious access to water for agriculture are major constraints to improving living conditions (Castillo et al., 2007). Water appears to be the most limiting factor for plant production. In arid and semi-arid areas, water losses through evapotranspiration are extremely high due to high temperature, sunshine and wind (Faurès et al., 2008). Thus, the exploration of water-efficient accessions is important for improving plant productivity in general and that of legumes in particular. Vigna subterranea (L.) can play an important role because one of its main characteristics is its tolerance to poor soils and drought, as well as its ability to produce under environmental stress conditions where other plants fail. However, Bambara groundnut yields remain low due to numerous production constraints. In the Far North region (Sahelian zone) of Cameroon, Bambara groundnut is generally grown on rocky soils, yet it has ecological elasticity (Basu et al., 2007). These types of soils are very porous and the plants very often face water stress. Thus, climate variability and irregular rainfall could intensify yield declines of Bambara groundnut in the Far North of Cameroon, marked by a contrasting climate. In order to overcome this shortcoming, a number of researchers have carried out extensive research on Bambara groundnut. As demonstrated by Ibrahima et al. (2024) in their study on the selection strategy for early maturing accessions capable of optimizing water uptake and use efficiency and Temegne et al. (2018) in their research on the combination of water deficit and phosphate fertilizer. It would therefore be important to explore the least water-demanding accessions. Thus, to contribute to the production of Bambara groundnut accessions that are physiologically adapting to water deficits, an assessment of the morphological and physiological characteristics of these accessions is necessary.
Field experiments were conducted at two sites: an experimental station (Site A: 10o33'N, 14o16'E) and a selected site (Site B: 10o32'N, 14o14'E). These sites are located in the Maroua 1^(er) district in the Far North region of Cameroon. They are characterized by a Sudano-Sahelian climate, with average temperatures between 28oC and 29oC, cumulative annual rainfall of 700 mm and savannah vegetation with sandy, calcareous, poorly evolved Vertisols and tropical ferruginous soils (Orstom, 1965). Seeds of four Bambara groundnut accessions were used as the initial genetic material. Four accessions were used for the pot and field experiments. They were chosen by phenotypic selection based on tegument color (Table 1).

Table 1: Characteristics of accessions used and their origins.


       
The trial was set up using a completely randomized block design with four replications. The seeds of the four cultivars were randomly sown by block at the experimental station and in the field at the Institute for Agricultural Research and Development (IRAD) in Maroua-Cameroon. Trials at different sites were carried out from 2022 to 2023. The pot experiment was carried out during the May- October 2022 period, while the field experiment was carried out during the June-November 2023 period. Water stress was induced for these accessions 15 days after sowing. The plants were subjected to three levels of water stress: 1 L, 0.5 L and 0.25 L of water per pot every five days for the first, second and third stress levels, respectively (in-station setup). Plants in the field setup were watered every 12 hours (morning and evening).
       
The experiment was conducted in to experimental sites in order to evaluate  the effect of water stress on Bambara groundnut accessions and assess the physiological traits such as relative water content (TRE), water loss rate (TDE), transpiration (TP), number of stomata (NS), leaf area, water use efficiency (WUE) and stress tolerance index (Table 2). The following morphological traits ere also evaluated: Germination time, germination rate, pod weight, seed weight, leaf length, leaf width, number of leaves, plant height, dry biomass, number of pods, number of seeds and grain yield (Table 2).

Table 2: Morphological, Physiological traits measured.


       
A statistical analysis of the variables was performed using SPSS.20 software. The collected data were then subjected to a series of statistical analyses, including an analysis of variance (ANOVA) and a Fisher test, which were employed to compare the means of the data sets. This test has facilitated the identification of varieties that exhibit substantial differences from one another. The Pearson correlation test was performed using XLSTAT version 2025.1 software.
Variation in morphological traits of accessions
 
Variation in germination time
 
The statistical outcomes presented in Table 3 demonstrate the variation in germination time among the diverse accessions. The variation in germination time was found to be significantly different between accessions (p = 0.000) and not between blocks (p = 0.42). Furthermore, the interaction between accessions and blocks was also found to be significant. Consequently, the blocks do not exert an influence on the germination time. This outcome is consistent with the hypothesis that the four blocks were exposed to identical conditions.

Table 3: Variation in germination time of Bambara groundnut accessions.


       
However, the accession V1 exhibited accelerated germination, occurring six days after sowing. The process of germination of the accession V4 is observed to occur at a later stage than that of the white accession, with a delay of three days in the onset of this process. The remaining two are considered intermediate. In all the accessions studied, radicle emergence occurred between six and nine days after sowing. The result obtained is analogous to that previously reported by Wassouo et al. (2019), who documented a germination time range of 6-7 days for Bambara groundnut. The findings of this study demonstrate a congruence with those of Touré et al. (2012), who reported that the germination of seeds in Bambara groundnut occurs within the timeframe of 6 to 15 days following the process of sowing. Karikari (1999) observed significantly higher values (14 to 24). The observed variability can be attributed to the presence of diverse plant material, variable climatic conditions and the varying quality of the seeds.
 
Variation in germination rate
 
A subsequent analysis of the results indicates a significant difference (Fig 1) between the accessions in terms of germination rate (p = 0.006). However, accession V1 exhibited the lowest germination rate (44%), while accession V2 demonstrated the highest (88%). However, no significant differences were observed between the V1 and V4 accession, the V4 and V3, or the V3 and V2. This negligible discrepancy may be attributable to the substantial variation observed among individuals (i.e. high standard deviations), given the limited understanding of the origins of these seeds and their parents. However, the mean germination rate was 66%. This value is higher than that reported by Touré et al. (2013), who found that in Bambara peanut accessions, the average germination rate is 58% at 30 days after sowing. Accession V2 exhibited the highest (88%). Consequently, it can be deduced from this observation that accessions exhibiting high germination rates would possess sufficient plants to ensure high yields, thereby conferring them with a certain agronomic interest in enhancing Bambara groundnut cultivation in the Sahelian regions.

Fig 1: Variation in germination rate according to accession.


 
Variation in length
 
The statistical analysis performed demonstrated that leaf length exhibited significant variation between accessions (p = 0.000) (Fig 2 and 4). However, the leaves of accession V1 are notably longer (7 cm) than those of the other three accessions. The shortest of these are the accession V3, which measure 5.5 centimetres. These results are identical to those of (Djè et al., 2005) who reported in Côte d’Ivoire that morphotypes of Bambara groundnut (White, Black and Red), the white variety gave the longest leaflets than the red and black varieties. Similar to the results of Bonny et al. (2011) for white and brown varieties, which had the longest leaflets compared to the others.

Fig 2: Variation in Length according to accessions.


 
Variation in width
 
The results of the analysis demonstrate the presence of variability in leaf widths (Fig 3). Indeed, a significant variation in leaf width was observed between accessions (p = 0.000), However, accession V1 exhibited the widest leaflets (2.8 cm), which was greater than the other accessions. These results differ from those reported by Yao et al. (2005), who found that the in Côte d’Ivoire, the Rouge variety was found to have the widest leaflets. than the white and black varieties. The leaf width observed in accession V1 indicates that these plants possess extensive photosynthetic surfaces, a trait that is likely attributable to the substantial light absorption capacity of these leaves in the Sahelian region, where temperatures are notably high. Consequently, they accumulate more organic matter.

Fig 3: Variation in Width according to accessions.



Fig 4: Pictorial presentation of the morphology of the plant at different growth stages.


       
The results from the descriptive statistical analysis (Table 4) demonstrate that, the mean number of leaves was determined to be seven at 40 days. This finding is incongruent with the observations reported by Wassouo et al. (2019), who documented that the mean number of leaves in Bambara groundnut is equivalent to 40 at 56 days post-sowing. The observed discrepancy can be attributed to the temporal increase in the average number of leaves. The mean plant height was found to be 15.08 centimeters, which is different to the findings of Temegne et al. (2018) who reported that the mean height of Bambara groundnut plants was 31,18 centimeters at 56 days post-sowing. This slight difference can be attributed to the fact that the growth of Bambara groundnut plants stabilizes after a certain number of days following the initial sowing. Consequently, the height of the plant exhibits a range from 9 centimeters to 20 centimeters. The values obtained in this study are analogous to those reported by Touré et al. (2013) who found values of 14 centimeters. However, these values are lower than those reported by Ndiang et al., (2012) in Cameroon, who found values ranging from 20 to 38 centimeters. The observed differences can be attributed to the genotype of each plant material, which is susceptible to environmental influences. A study of Bambara groundnut accessions on quantitative traits was reported by Khan et al., (2020), who noted a coefficient of variation (CV) ≥20% for quantitative traits studied. This finding aligns with the results of our study on variation coefficients, suggesting heterogeneity within the plant material for these variables. A similar, relevant observation of a high coefficient of variation was confirmed by Goli et al. (1997).

Table 4: Descriptive statistics on morphological parameters.


 
Variation in yield components and dry matter (Table 5 and 6)

Table 5: Variation of yield components and dry matter.



Table 6: Effect of water stress on yield components and dry matter.


 
A thorough examination of the data reveals a statistically significant impact of water stress on the overall weight of 100 pods and seeds produced per accession. Accession V1 exhibited a significant response to water stress, resulting in the production of the smallest pods (22 g). These results are consistent with those reported by Erskine et al. (1993), who found that abiotic stress significantly reduces lentil yield. In contrast, accession V4 exhibited a reduced impact, yielding an average pod weight of 24 grams. Accessions V2 and V3 exhibited the highest recorded weight of 26 g. These two accessions can be regarded as exhibiting reduced sensitivity to water stress. Consistent findings have been reported in related studies (Tambal et al., 2000). On average, P100Gss exhibited a discrepancy of approximately 0.6 g in favor of the V3 accession. This phenomenon does not appear to exert a substantial influence on yield in our experimental setting, as evidenced by the observation that accession V2, characterized by a low pod weight of 26.14 g, exhibits the highest seed weight of 24.99 g. The total number of seeds is diminished; however, the reduction in the number of sheaths due to water stress is comparatively greater in accession V1, which has seeds of a lighter hue. The range of these values was from 6.16 in accession V1 to 9.47 in accession V2. Water stress has been demonstrated to result in pod abortion, which is characterized by a reduction in seed number due to the loss of young pods prior to seed formation and filling. These results are consistent with those reported by Shrestha et al. (2006), who observed that water stress led to a 27% reduction in the number of pods, with a significantly higher decrease in empty pods when water was retained.  However, a higher root biomass weight was observed in accession V1 compared to the other accessions. However, this variability among accessions can be attributed to the observation that early accessions consistently yield high levels of dry matter. The findings of this study are consistent with those reported by Ouedraogo ​et al. (2008), who observed that late varieties yielded the least. The range of above-ground biomass values was from 5.34 grams in accession V1 to 4.19 grams in V2. Accession V1 exhibited the highest above-ground biomass value in comparison with accession V2. Despite its notably high above-ground dry weight, accession V1 exhibited the greatest sensitivity to water stress among the accessions examined, resulting in the lowest grain yield. This phenomenon could be attributed to two potential factors. Firstly, there is the possibility that vegetative organisms are being disproportionately assimilated under moderate conditions. Alternatively, the stressed V1 accession may have allocated a greater proportion of its resources to biomass growth, thereby compromising grain yield. The present findings contradict those of Bouzerzour et al. (1998), who posited that in variable environments, ensuring sufficient above-ground biomass production is imperative to ensure an acceptable yield. However, similar results were observed in the study by Surson et al. (2025), who reported that the rice M2 RD.43 variant 1 had more leaves but fewer whole seeds, as well as lower whole seed weight, seed width, seed length and seed formation percentage.

Variation of physiological traits of accessions
 
A statistical analysis of Table 7 reveals that the maximum error ratio (TRE) was observed at N1 (1L/5D), while the minimum was observed at N3 (0.25/5D). This finding suggests that the TRE declines with increasing water deficit. These results are consistent with those reported by Albouchi et al. (2000), who observed that the TRE in durum wheat declines as soil water content decreases. Lilya et al. (2002) observed that accessions capable of maintaining a high TRE in the presence of water stress are classified as tolerant accessions. A minimum water loss (TDE) of (-2.114± 0.067a) was observed in accession V2 at a severe stress level (N3, 0.25L 5D) compared with a high-water loss (-19.721±0.063c) recorded in accession V1. Additionally, accession V2 exhibited the smallest leaf area at (N3, 0.25L/5d). This suggests that the rate of water loss is inversely proportional to the surface area, with smaller surface areas resulting in reduced water loss. The results of this study are consistent with those reported by Kelliher et al. (1980). The aforementioned researchers observed that, in durum wheat, the TDE increases as the leaf area expands. Accession V1 exhibited the largest surface area across all stress levels, while accession V2 demonstrated the smallest surface area in comparison to the other accessions. The smallest leaf areas were recorded at the severe stress level (N3, 0.25 L/5 d).

Table 7: Influence of the level of water stress on the physiological parameters.


       
This finding suggests a negative correlation between leaf area and water deficit, indicating that as the water deficit increases, leaf area decreases. According to Granier et al. (2000), the leaves of plants subjected to water deficit exhibited smaller apparent final sizes in comparison to the control group. Lebon et al. (2006) confirmed that one of the earliest responses of plants to water deficit is the reduction of leaf area. Lebon et al. (2006) demonstrated that the reduction in leaf area under water is an adaptive mechanism for plants to limit leaf transpiration when water conditions become unfavorable. Conversely, only accessions V3 and V2 exhibited a diminished number of stomata among the accessions examined. A substantial decrease in the quantity of stomata was detected under conditions of severe stress (N3, 0.25 L/5 d) across all accessions, with the water deficit leading to a reduction of approximately 10 to 20% in stomata, there by constraining transpiration. This is explained by the fact that the onset of water stress implies a reduction in water loss, through the closure of stomata, resulting in a reduction in photosynthetic activity and transpiration. Thus, accession V2 exhibited a reduced level of transpiration in comparison to the other accessions. The findings of this study demonstrate a congruence with the results previously reported by Bourou et al. (2022) on the subject of Tamarindus, wherein a correlation was established between transpiration, stomatal conductance and net photosynthesis. A low water use efficiency (WUE) was observed (Fig 3) in accession V1 (0.80g/l) in contrast to accession V2, which has a higher WUE (0.89g/l). Accessions V3 and V4 demonstrate intermediate WUE values. The mean WUE values obtained from the accessions range from 0.80 g/l to 0.89 g/l. The findings of Chetto et al. (2020) indicated an average value of 0.712 WUE for the farms under analysis, signifying an average of 71.2% of the WUE by citrus growers. This finding aligns with the conclusions drawn from the present study. These divergent outcomes are consistent with the findings reported by Karam et al. (2007), who conducted a study on the water use efficiency (WUE) in corn. This phenomenon can be attributed to the higher WUE exhibited by plants with C4 photosynthetic metabolism in comparison to C3 species (Rubino et al., 1999). The V2 model demonstrated a high level of water use efficiency, as illustrated in Fig 5. This phenomenon can be attributed to the established relationship between yield and WUE, wherein elevated levels of yield are accompanied by heightened WUE. Therefore, it can be concluded from this observation that increases in yield are associated with accessions that demonstrate effective water use efficiency.

Fig 5: Variation of WUE with respect to accessions.


       
Table 8 presents the characteristics of accessions on yield in real conditions and under stress In the second season, an experiment was conducted at two distinct locations with the objective of estimating the mean yield per accession. The accessions were evaluated under two distinct conditions: Water-limited and natural. An evaluation of the test results indicates that accession V2 exhibits a significantly higher grain yield (65.72±0.50b) under conditions of water limitation in comparison to the other accessions. These results are similar to those of Karthiga et al. (2025), who observed an increase in cucumber productivity when genotypes were evaluated under water-deficient conditions. These results are also in line with observations of (Dolinassou et al., 2016) on Arrachis hypogea, who reported that in the same location, the Gobo-55-437 line exhibited the highest average yield, while the NW-Red Esimbi genotype demonstrated a low yield.Water deficits have been shown to reduce yield by up to 10% in the V1 accession. Nevertheless, accession V3 demonstrated the highest yield (65.50±0.47 g) when the accessions were assessed in the field. These results are consistent with those reported by Shrestha et al. (2006), who found that seed yield in Lens culinaris was significantly higher than in other genotypes under conditions of adequate irrigation. Among the accessions examined, only two (accession V3 and V2) exhibited the optimal yield under stress conditions, with values of 64.96 g and 65.72 g, respectively. The observed outcomes can be attributed to the combined effects of water shock and end-of-season drought in this agroecological zone. However, it is noteworthy that only the V1 accession exhibited a reduction in grain yield. The decline in grain yield under water stress is predominantly attributed to the reduction in the number of seeds. These results are consistent with the observations of Shrestha et al. (2006), who noted that in Lens culinaris medikus, the 70% reduction in seed yield induced by water deficit was primarily due to a decrease in the number of pods and seeds. The utilization of the agronomic trait (number of pods and seeds) can facilitate the identification of accessions with superior traits (Gahoonia et al., 2006) that can be integrated into the breeding program for the production of drought-tolerant varieties.

Table 8: Yield variability in real conditions and under stress.


       
The highest stress tolerance indices were observed in accessions V2 and V3, while accession V1 was the most sensitive (Table 7). Similar results reported by Mohammadi et al. (2011) indicated that the index (STI) is well suited for selecting the most productive RILs (Recombinant Inbred Lines) under both stressful and favorable conditions. It is therefore clear that the stress tolerance index (STI) is a strong discriminator between genotypes with high yields and potential tolerance to water stress and other genotypes. According to Fernandez (1992), the STI can be used to identify genotypes that produce high yields under both favorable and stressful conditions.
 
Correlation matrix between the parameters studied
 
Table 9 presents the Pearson correlation matrix between the traits studied. In the genetic variability study of accessions by Khan et al., 2020, it was concluded that a strong positive correlation (0.75 <r <1) was found for the seed weight and pod weight traits. A similar variation in these traits was reported by Goli et al. (1995), which confirms our results on the observation of strong and significant correlations between the weight of 100 seeds and the weight of 100 pods (r=0.956; p<0.01). This measurement of the weight of 100 seeds was considered an essential tool for evaluating morpho-physiological traits related to yield (Mohammed et al., 2014). Kirham et al., 1980 noted that in durum wheat, the TDE increases as the surface area expands. A strong correlation was observed between germination time and leaf length (r = 0.736, p<0.01). Thus, the earliest accessions have the longest and widest leaves, indicating that the earlier they germinate, the more organic matter they accumulate. Additionally, the larger the leaf surface area, the higher the growth rate and yield. Cherfia et al. (2010) demonstrate a significant relationship between leaf area and yield. Indeed, the greater the leaf area, the higher the yield. Patel et al. (2025) found that larger leaflets can exploit more sunlight, resulting in greater biomass accumulation and higher yields. Araus et al., (1998) report that selecting for large leaf area improves grain yields. Belkharchouche et al. (2009) report that leaf area determines drought resistance since high leaf areas lose more water than low leaf areas.

Table 9: Correlation matrix between the parameters studied.

The present study was conducted on Bambara groundnut accessions with the objective of evaluating the morphological and physiological characteristics of this species. The results demonstrated significant variability among the accessions under water stress. In regard to physiological parameters, the impact of water stress proved to be highly significant across the majority of the physiological traits that were examined. Water stress was applied to the accessions to identify an accession least sensitive to water stress. Accession V2 was identified as the most resistant, as indicated by lower water loss values, which are indicative of reduced sensitivity to stress. The diverse morphological responses exhibited by the accessions’ genetic material indicated that accession V2 demonstrated the least significant impact, exhibiting the highest 100-pod weight of 26 grams. This is in contrast to accession V1, which exhibited a substantial response to water stress, resulting in the lowest pod weight recorded at 22 grams. However, accession V2 exhibited a substantially higher seed weight compared to the other accessions. Given its notable productivity under conditions of hydric stress, this V2 accession has the potential to be selected for the enhancement of Bambara groundnut production in the Sahelian regions of Cameroon. However, further investigation into the water stress effect at other levels in the field with these accessions could facilitate the completion of these results for their use in breeding and varietal improvement programs.
The authors are thankful for the bench space and technical support from Bio-Lab Foods Company Ltd during laboratory experiments.
 
Disclaimers
 
The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.
 
Informed consent
 
Not applicable.
The authors declare no conflicts of interest.

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Effect of Water Stress on Morpho-physiological Trait of Some Accessions of Bambara Groundnut [Vigna subterranea (L). Verdcourt]

A
Abib Chimene Fanta1
W
Wassouo Alain Felix2
N
Nassourou Maina Antoine1
D
Difo Voukang Harouna1
T
Tchapda Dorothy Tchatchoua2
1Department of Biological Sciences, Faculty of Sciences, University of Maroua, Po. Box. 814 Maroua, Cameroon.
2Department of Agriculture, Animal Husbandry and Derived Products, National Advanced School of Engineering, University of Maroua. Po. Box. 46 Maroua, Cameroon.
  • Submitted21-05-2025|

  • Accepted29-07-2025|

  • First Online 15-09-2025|

  • doi 10.18805/LRF-878

Background: The cultivation of Bambara groundnut in arid zones, such as the Sahelian regions of Cameroon, has received scant attention despite its nutritional and agronomic potential. The Sahelian regions of Cameroon are classified as part of the arid zones of the world, characterized by a rainy season of approximately three to four months in duration. Nevertheless, the cultivation period of Bambara groundnut is approximately four to six months, a duration that hinders its cultivation. The objective of this study is to assess the impact of water stress on the morpho-physiological traits of select Bambara groundnut accessions cultivated in the Sahelian regions of Cameroon.

Methods: The experiment was executed in accordance with the fisher bloc design, employing four replicates. The water stress was induced using distilled water in pot experiments to monitor thirteen (13) morphological traits and six (06) physiological traits.

Result: The results demonstrated significant variability among the accessions under water stress. Accession V2 exhibited the greatest resistance, characterized by diminished leaf areas and reduced stomatal number. The findings demonstrated a consistent increase in relative water content, accompanied by a decrease in water loss and transpiration. This observation suggests a reduced response to stress. The genetic material of the accession exhibited a range of morphological responses. The considerable impact of water stress was evident on the 100-pod weight produced per accession. Accession V1 exhibited a significant response to water stress, resulting in the production of the smallest pods (22 g). Accession V2 demonstrated the least susceptibility to the 100-pod weight (26 g), exhibiting the highest weight among the other accessions. The evaluation of the 100 seeds’ weight revealed that accession V2 exhibited a significantly higher seed weight compared to the other accessions. The grain yield in the accession V2 was higher for both sites (65.39±1.15 and 65.72±0.50). Principal component analysis demonstrated a positive correlation between stress and the weights of 100 pods and seeds (r = 0.956). The results obtained underscore the capacity of accession V2 to adapt to the environmental conditions prevalent in the Sahelian regions of Cameroon.

Global population growth over the past few decades is generating enormous needs, the most notable of which in sub-Saharan Africa are energy and food demand. in fact, the report published by the FAO, shows the absolute number of people suffering from hunger and under nourishment reached almost 821 million in 2017, compared with around 804 million in 2016, (Salomon et al., 2023), Indeed, in developing countries, 12.9% of the population is undernourished and malnutrition is the cause of 45% of deaths in children under 5.
       
[Vigna subterranea (L.) Verct], commonly known as Bambara groundnut, is a plant indigenous to Africa and is one of the often-neglected subsistence crops in Africa (Soumare et al., 2022). It is a minor food legume adapted to a variety of stressful climatic and ecological conditions (Onwubiko et al., 2011). It is a calorie-rich plant food, with an energy value of around 387 Kcal/100 g. It is also rich in vitamins, minerals, proteins (20-25%), lipids (6-8%) and carbohydrates (50-57%) (Gbaguidi et al., 2015). Bambara peanut seeds are used as human food as well as for the treatment of certain pathogens (Gbaguidi et al., 2015). However, many plant species grown around the world for food production are considered minor crops compared to other food crops considered major food crops (rice, maize, wheat, sorghum, yam, plantain, cassava, etc.) which make up almost the entire diet in Africa (Bricas et al., 2009). However, these minor crops play a crucial role in food security, nutrition and generate income for many rural populations (Ouoba et al., 2018). Despite its nutritional qualities, Bambara groundnut is on the list of neglected species in varietal breeding programs (Tadele Zerihun, 2009). However, the plant offers enormous potential for improvement, thanks to its genetic diversity found in thousands of accessions harvested throughout the world. (Djè et al., 2005); (Ndiang et al., 2012). In Cameroon for instance, very little information is currently available on the genetic and morpho-physiological diversity of this species.
               
In the arid regions of Cameroon, water scarcity and precarious access to water for agriculture are major constraints to improving living conditions (Castillo et al., 2007). Water appears to be the most limiting factor for plant production. In arid and semi-arid areas, water losses through evapotranspiration are extremely high due to high temperature, sunshine and wind (Faurès et al., 2008). Thus, the exploration of water-efficient accessions is important for improving plant productivity in general and that of legumes in particular. Vigna subterranea (L.) can play an important role because one of its main characteristics is its tolerance to poor soils and drought, as well as its ability to produce under environmental stress conditions where other plants fail. However, Bambara groundnut yields remain low due to numerous production constraints. In the Far North region (Sahelian zone) of Cameroon, Bambara groundnut is generally grown on rocky soils, yet it has ecological elasticity (Basu et al., 2007). These types of soils are very porous and the plants very often face water stress. Thus, climate variability and irregular rainfall could intensify yield declines of Bambara groundnut in the Far North of Cameroon, marked by a contrasting climate. In order to overcome this shortcoming, a number of researchers have carried out extensive research on Bambara groundnut. As demonstrated by Ibrahima et al. (2024) in their study on the selection strategy for early maturing accessions capable of optimizing water uptake and use efficiency and Temegne et al. (2018) in their research on the combination of water deficit and phosphate fertilizer. It would therefore be important to explore the least water-demanding accessions. Thus, to contribute to the production of Bambara groundnut accessions that are physiologically adapting to water deficits, an assessment of the morphological and physiological characteristics of these accessions is necessary.
Field experiments were conducted at two sites: an experimental station (Site A: 10o33'N, 14o16'E) and a selected site (Site B: 10o32'N, 14o14'E). These sites are located in the Maroua 1^(er) district in the Far North region of Cameroon. They are characterized by a Sudano-Sahelian climate, with average temperatures between 28oC and 29oC, cumulative annual rainfall of 700 mm and savannah vegetation with sandy, calcareous, poorly evolved Vertisols and tropical ferruginous soils (Orstom, 1965). Seeds of four Bambara groundnut accessions were used as the initial genetic material. Four accessions were used for the pot and field experiments. They were chosen by phenotypic selection based on tegument color (Table 1).

Table 1: Characteristics of accessions used and their origins.


       
The trial was set up using a completely randomized block design with four replications. The seeds of the four cultivars were randomly sown by block at the experimental station and in the field at the Institute for Agricultural Research and Development (IRAD) in Maroua-Cameroon. Trials at different sites were carried out from 2022 to 2023. The pot experiment was carried out during the May- October 2022 period, while the field experiment was carried out during the June-November 2023 period. Water stress was induced for these accessions 15 days after sowing. The plants were subjected to three levels of water stress: 1 L, 0.5 L and 0.25 L of water per pot every five days for the first, second and third stress levels, respectively (in-station setup). Plants in the field setup were watered every 12 hours (morning and evening).
       
The experiment was conducted in to experimental sites in order to evaluate  the effect of water stress on Bambara groundnut accessions and assess the physiological traits such as relative water content (TRE), water loss rate (TDE), transpiration (TP), number of stomata (NS), leaf area, water use efficiency (WUE) and stress tolerance index (Table 2). The following morphological traits ere also evaluated: Germination time, germination rate, pod weight, seed weight, leaf length, leaf width, number of leaves, plant height, dry biomass, number of pods, number of seeds and grain yield (Table 2).

Table 2: Morphological, Physiological traits measured.


       
A statistical analysis of the variables was performed using SPSS.20 software. The collected data were then subjected to a series of statistical analyses, including an analysis of variance (ANOVA) and a Fisher test, which were employed to compare the means of the data sets. This test has facilitated the identification of varieties that exhibit substantial differences from one another. The Pearson correlation test was performed using XLSTAT version 2025.1 software.
Variation in morphological traits of accessions
 
Variation in germination time
 
The statistical outcomes presented in Table 3 demonstrate the variation in germination time among the diverse accessions. The variation in germination time was found to be significantly different between accessions (p = 0.000) and not between blocks (p = 0.42). Furthermore, the interaction between accessions and blocks was also found to be significant. Consequently, the blocks do not exert an influence on the germination time. This outcome is consistent with the hypothesis that the four blocks were exposed to identical conditions.

Table 3: Variation in germination time of Bambara groundnut accessions.


       
However, the accession V1 exhibited accelerated germination, occurring six days after sowing. The process of germination of the accession V4 is observed to occur at a later stage than that of the white accession, with a delay of three days in the onset of this process. The remaining two are considered intermediate. In all the accessions studied, radicle emergence occurred between six and nine days after sowing. The result obtained is analogous to that previously reported by Wassouo et al. (2019), who documented a germination time range of 6-7 days for Bambara groundnut. The findings of this study demonstrate a congruence with those of Touré et al. (2012), who reported that the germination of seeds in Bambara groundnut occurs within the timeframe of 6 to 15 days following the process of sowing. Karikari (1999) observed significantly higher values (14 to 24). The observed variability can be attributed to the presence of diverse plant material, variable climatic conditions and the varying quality of the seeds.
 
Variation in germination rate
 
A subsequent analysis of the results indicates a significant difference (Fig 1) between the accessions in terms of germination rate (p = 0.006). However, accession V1 exhibited the lowest germination rate (44%), while accession V2 demonstrated the highest (88%). However, no significant differences were observed between the V1 and V4 accession, the V4 and V3, or the V3 and V2. This negligible discrepancy may be attributable to the substantial variation observed among individuals (i.e. high standard deviations), given the limited understanding of the origins of these seeds and their parents. However, the mean germination rate was 66%. This value is higher than that reported by Touré et al. (2013), who found that in Bambara peanut accessions, the average germination rate is 58% at 30 days after sowing. Accession V2 exhibited the highest (88%). Consequently, it can be deduced from this observation that accessions exhibiting high germination rates would possess sufficient plants to ensure high yields, thereby conferring them with a certain agronomic interest in enhancing Bambara groundnut cultivation in the Sahelian regions.

Fig 1: Variation in germination rate according to accession.


 
Variation in length
 
The statistical analysis performed demonstrated that leaf length exhibited significant variation between accessions (p = 0.000) (Fig 2 and 4). However, the leaves of accession V1 are notably longer (7 cm) than those of the other three accessions. The shortest of these are the accession V3, which measure 5.5 centimetres. These results are identical to those of (Djè et al., 2005) who reported in Côte d’Ivoire that morphotypes of Bambara groundnut (White, Black and Red), the white variety gave the longest leaflets than the red and black varieties. Similar to the results of Bonny et al. (2011) for white and brown varieties, which had the longest leaflets compared to the others.

Fig 2: Variation in Length according to accessions.


 
Variation in width
 
The results of the analysis demonstrate the presence of variability in leaf widths (Fig 3). Indeed, a significant variation in leaf width was observed between accessions (p = 0.000), However, accession V1 exhibited the widest leaflets (2.8 cm), which was greater than the other accessions. These results differ from those reported by Yao et al. (2005), who found that the in Côte d’Ivoire, the Rouge variety was found to have the widest leaflets. than the white and black varieties. The leaf width observed in accession V1 indicates that these plants possess extensive photosynthetic surfaces, a trait that is likely attributable to the substantial light absorption capacity of these leaves in the Sahelian region, where temperatures are notably high. Consequently, they accumulate more organic matter.

Fig 3: Variation in Width according to accessions.



Fig 4: Pictorial presentation of the morphology of the plant at different growth stages.


       
The results from the descriptive statistical analysis (Table 4) demonstrate that, the mean number of leaves was determined to be seven at 40 days. This finding is incongruent with the observations reported by Wassouo et al. (2019), who documented that the mean number of leaves in Bambara groundnut is equivalent to 40 at 56 days post-sowing. The observed discrepancy can be attributed to the temporal increase in the average number of leaves. The mean plant height was found to be 15.08 centimeters, which is different to the findings of Temegne et al. (2018) who reported that the mean height of Bambara groundnut plants was 31,18 centimeters at 56 days post-sowing. This slight difference can be attributed to the fact that the growth of Bambara groundnut plants stabilizes after a certain number of days following the initial sowing. Consequently, the height of the plant exhibits a range from 9 centimeters to 20 centimeters. The values obtained in this study are analogous to those reported by Touré et al. (2013) who found values of 14 centimeters. However, these values are lower than those reported by Ndiang et al., (2012) in Cameroon, who found values ranging from 20 to 38 centimeters. The observed differences can be attributed to the genotype of each plant material, which is susceptible to environmental influences. A study of Bambara groundnut accessions on quantitative traits was reported by Khan et al., (2020), who noted a coefficient of variation (CV) ≥20% for quantitative traits studied. This finding aligns with the results of our study on variation coefficients, suggesting heterogeneity within the plant material for these variables. A similar, relevant observation of a high coefficient of variation was confirmed by Goli et al. (1997).

Table 4: Descriptive statistics on morphological parameters.


 
Variation in yield components and dry matter (Table 5 and 6)

Table 5: Variation of yield components and dry matter.



Table 6: Effect of water stress on yield components and dry matter.


 
A thorough examination of the data reveals a statistically significant impact of water stress on the overall weight of 100 pods and seeds produced per accession. Accession V1 exhibited a significant response to water stress, resulting in the production of the smallest pods (22 g). These results are consistent with those reported by Erskine et al. (1993), who found that abiotic stress significantly reduces lentil yield. In contrast, accession V4 exhibited a reduced impact, yielding an average pod weight of 24 grams. Accessions V2 and V3 exhibited the highest recorded weight of 26 g. These two accessions can be regarded as exhibiting reduced sensitivity to water stress. Consistent findings have been reported in related studies (Tambal et al., 2000). On average, P100Gss exhibited a discrepancy of approximately 0.6 g in favor of the V3 accession. This phenomenon does not appear to exert a substantial influence on yield in our experimental setting, as evidenced by the observation that accession V2, characterized by a low pod weight of 26.14 g, exhibits the highest seed weight of 24.99 g. The total number of seeds is diminished; however, the reduction in the number of sheaths due to water stress is comparatively greater in accession V1, which has seeds of a lighter hue. The range of these values was from 6.16 in accession V1 to 9.47 in accession V2. Water stress has been demonstrated to result in pod abortion, which is characterized by a reduction in seed number due to the loss of young pods prior to seed formation and filling. These results are consistent with those reported by Shrestha et al. (2006), who observed that water stress led to a 27% reduction in the number of pods, with a significantly higher decrease in empty pods when water was retained.  However, a higher root biomass weight was observed in accession V1 compared to the other accessions. However, this variability among accessions can be attributed to the observation that early accessions consistently yield high levels of dry matter. The findings of this study are consistent with those reported by Ouedraogo ​et al. (2008), who observed that late varieties yielded the least. The range of above-ground biomass values was from 5.34 grams in accession V1 to 4.19 grams in V2. Accession V1 exhibited the highest above-ground biomass value in comparison with accession V2. Despite its notably high above-ground dry weight, accession V1 exhibited the greatest sensitivity to water stress among the accessions examined, resulting in the lowest grain yield. This phenomenon could be attributed to two potential factors. Firstly, there is the possibility that vegetative organisms are being disproportionately assimilated under moderate conditions. Alternatively, the stressed V1 accession may have allocated a greater proportion of its resources to biomass growth, thereby compromising grain yield. The present findings contradict those of Bouzerzour et al. (1998), who posited that in variable environments, ensuring sufficient above-ground biomass production is imperative to ensure an acceptable yield. However, similar results were observed in the study by Surson et al. (2025), who reported that the rice M2 RD.43 variant 1 had more leaves but fewer whole seeds, as well as lower whole seed weight, seed width, seed length and seed formation percentage.

Variation of physiological traits of accessions
 
A statistical analysis of Table 7 reveals that the maximum error ratio (TRE) was observed at N1 (1L/5D), while the minimum was observed at N3 (0.25/5D). This finding suggests that the TRE declines with increasing water deficit. These results are consistent with those reported by Albouchi et al. (2000), who observed that the TRE in durum wheat declines as soil water content decreases. Lilya et al. (2002) observed that accessions capable of maintaining a high TRE in the presence of water stress are classified as tolerant accessions. A minimum water loss (TDE) of (-2.114± 0.067a) was observed in accession V2 at a severe stress level (N3, 0.25L 5D) compared with a high-water loss (-19.721±0.063c) recorded in accession V1. Additionally, accession V2 exhibited the smallest leaf area at (N3, 0.25L/5d). This suggests that the rate of water loss is inversely proportional to the surface area, with smaller surface areas resulting in reduced water loss. The results of this study are consistent with those reported by Kelliher et al. (1980). The aforementioned researchers observed that, in durum wheat, the TDE increases as the leaf area expands. Accession V1 exhibited the largest surface area across all stress levels, while accession V2 demonstrated the smallest surface area in comparison to the other accessions. The smallest leaf areas were recorded at the severe stress level (N3, 0.25 L/5 d).

Table 7: Influence of the level of water stress on the physiological parameters.


       
This finding suggests a negative correlation between leaf area and water deficit, indicating that as the water deficit increases, leaf area decreases. According to Granier et al. (2000), the leaves of plants subjected to water deficit exhibited smaller apparent final sizes in comparison to the control group. Lebon et al. (2006) confirmed that one of the earliest responses of plants to water deficit is the reduction of leaf area. Lebon et al. (2006) demonstrated that the reduction in leaf area under water is an adaptive mechanism for plants to limit leaf transpiration when water conditions become unfavorable. Conversely, only accessions V3 and V2 exhibited a diminished number of stomata among the accessions examined. A substantial decrease in the quantity of stomata was detected under conditions of severe stress (N3, 0.25 L/5 d) across all accessions, with the water deficit leading to a reduction of approximately 10 to 20% in stomata, there by constraining transpiration. This is explained by the fact that the onset of water stress implies a reduction in water loss, through the closure of stomata, resulting in a reduction in photosynthetic activity and transpiration. Thus, accession V2 exhibited a reduced level of transpiration in comparison to the other accessions. The findings of this study demonstrate a congruence with the results previously reported by Bourou et al. (2022) on the subject of Tamarindus, wherein a correlation was established between transpiration, stomatal conductance and net photosynthesis. A low water use efficiency (WUE) was observed (Fig 3) in accession V1 (0.80g/l) in contrast to accession V2, which has a higher WUE (0.89g/l). Accessions V3 and V4 demonstrate intermediate WUE values. The mean WUE values obtained from the accessions range from 0.80 g/l to 0.89 g/l. The findings of Chetto et al. (2020) indicated an average value of 0.712 WUE for the farms under analysis, signifying an average of 71.2% of the WUE by citrus growers. This finding aligns with the conclusions drawn from the present study. These divergent outcomes are consistent with the findings reported by Karam et al. (2007), who conducted a study on the water use efficiency (WUE) in corn. This phenomenon can be attributed to the higher WUE exhibited by plants with C4 photosynthetic metabolism in comparison to C3 species (Rubino et al., 1999). The V2 model demonstrated a high level of water use efficiency, as illustrated in Fig 5. This phenomenon can be attributed to the established relationship between yield and WUE, wherein elevated levels of yield are accompanied by heightened WUE. Therefore, it can be concluded from this observation that increases in yield are associated with accessions that demonstrate effective water use efficiency.

Fig 5: Variation of WUE with respect to accessions.


       
Table 8 presents the characteristics of accessions on yield in real conditions and under stress In the second season, an experiment was conducted at two distinct locations with the objective of estimating the mean yield per accession. The accessions were evaluated under two distinct conditions: Water-limited and natural. An evaluation of the test results indicates that accession V2 exhibits a significantly higher grain yield (65.72±0.50b) under conditions of water limitation in comparison to the other accessions. These results are similar to those of Karthiga et al. (2025), who observed an increase in cucumber productivity when genotypes were evaluated under water-deficient conditions. These results are also in line with observations of (Dolinassou et al., 2016) on Arrachis hypogea, who reported that in the same location, the Gobo-55-437 line exhibited the highest average yield, while the NW-Red Esimbi genotype demonstrated a low yield.Water deficits have been shown to reduce yield by up to 10% in the V1 accession. Nevertheless, accession V3 demonstrated the highest yield (65.50±0.47 g) when the accessions were assessed in the field. These results are consistent with those reported by Shrestha et al. (2006), who found that seed yield in Lens culinaris was significantly higher than in other genotypes under conditions of adequate irrigation. Among the accessions examined, only two (accession V3 and V2) exhibited the optimal yield under stress conditions, with values of 64.96 g and 65.72 g, respectively. The observed outcomes can be attributed to the combined effects of water shock and end-of-season drought in this agroecological zone. However, it is noteworthy that only the V1 accession exhibited a reduction in grain yield. The decline in grain yield under water stress is predominantly attributed to the reduction in the number of seeds. These results are consistent with the observations of Shrestha et al. (2006), who noted that in Lens culinaris medikus, the 70% reduction in seed yield induced by water deficit was primarily due to a decrease in the number of pods and seeds. The utilization of the agronomic trait (number of pods and seeds) can facilitate the identification of accessions with superior traits (Gahoonia et al., 2006) that can be integrated into the breeding program for the production of drought-tolerant varieties.

Table 8: Yield variability in real conditions and under stress.


       
The highest stress tolerance indices were observed in accessions V2 and V3, while accession V1 was the most sensitive (Table 7). Similar results reported by Mohammadi et al. (2011) indicated that the index (STI) is well suited for selecting the most productive RILs (Recombinant Inbred Lines) under both stressful and favorable conditions. It is therefore clear that the stress tolerance index (STI) is a strong discriminator between genotypes with high yields and potential tolerance to water stress and other genotypes. According to Fernandez (1992), the STI can be used to identify genotypes that produce high yields under both favorable and stressful conditions.
 
Correlation matrix between the parameters studied
 
Table 9 presents the Pearson correlation matrix between the traits studied. In the genetic variability study of accessions by Khan et al., 2020, it was concluded that a strong positive correlation (0.75 <r <1) was found for the seed weight and pod weight traits. A similar variation in these traits was reported by Goli et al. (1995), which confirms our results on the observation of strong and significant correlations between the weight of 100 seeds and the weight of 100 pods (r=0.956; p<0.01). This measurement of the weight of 100 seeds was considered an essential tool for evaluating morpho-physiological traits related to yield (Mohammed et al., 2014). Kirham et al., 1980 noted that in durum wheat, the TDE increases as the surface area expands. A strong correlation was observed between germination time and leaf length (r = 0.736, p<0.01). Thus, the earliest accessions have the longest and widest leaves, indicating that the earlier they germinate, the more organic matter they accumulate. Additionally, the larger the leaf surface area, the higher the growth rate and yield. Cherfia et al. (2010) demonstrate a significant relationship between leaf area and yield. Indeed, the greater the leaf area, the higher the yield. Patel et al. (2025) found that larger leaflets can exploit more sunlight, resulting in greater biomass accumulation and higher yields. Araus et al., (1998) report that selecting for large leaf area improves grain yields. Belkharchouche et al. (2009) report that leaf area determines drought resistance since high leaf areas lose more water than low leaf areas.

Table 9: Correlation matrix between the parameters studied.

The present study was conducted on Bambara groundnut accessions with the objective of evaluating the morphological and physiological characteristics of this species. The results demonstrated significant variability among the accessions under water stress. In regard to physiological parameters, the impact of water stress proved to be highly significant across the majority of the physiological traits that were examined. Water stress was applied to the accessions to identify an accession least sensitive to water stress. Accession V2 was identified as the most resistant, as indicated by lower water loss values, which are indicative of reduced sensitivity to stress. The diverse morphological responses exhibited by the accessions’ genetic material indicated that accession V2 demonstrated the least significant impact, exhibiting the highest 100-pod weight of 26 grams. This is in contrast to accession V1, which exhibited a substantial response to water stress, resulting in the lowest pod weight recorded at 22 grams. However, accession V2 exhibited a substantially higher seed weight compared to the other accessions. Given its notable productivity under conditions of hydric stress, this V2 accession has the potential to be selected for the enhancement of Bambara groundnut production in the Sahelian regions of Cameroon. However, further investigation into the water stress effect at other levels in the field with these accessions could facilitate the completion of these results for their use in breeding and varietal improvement programs.
The authors are thankful for the bench space and technical support from Bio-Lab Foods Company Ltd during laboratory experiments.
 
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The views and conclusions expressed in this article are solely those of the authors and do not necessarily represent the views of their affiliated institutions. The authors are responsible for the accuracy and completeness of the information provided, but do not accept any liability for any direct or indirect losses resulting from the use of this content.
 
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The authors declare no conflicts of interest.

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