Variation of faba bean traits under irrigation regimes
Combined analysis of variance for faba bean traits (NTS, NP, NS, WSS and PY) under different water regimes is summarized in Table 1.
Across environments and genotypes, highly significant differences were detected for most traits, indicating both the environment and genetic background contributed appreciably to total variation (Table 1). For NTS and NP, the environment accounted for 8.65% and 33.40% of the total SS, respectively, with mean squares that were significant at
P<0.05 for NTS and at
P<0.001 for NP. This shows that both NTS and NP are substantially influenced by the irrigation regime. In contrast, the environment had little influence on NS (1.26%, not significant) and a moderate but highly significant effect on WSS (22.18%,
P<0.001) and PY (61.40%,
P<0.001), indicating that environment plays a critical role in determining WSS and especially PY. The variety (genotype) source of variation was highly significant (
P<0.001) for all five traits, explaining 12.40% of variation in NTS, 102.85% in NP, 154.20% in NS, 86.73% in WSS and 98.50% in PY. These large percentages reflect strong genetic divergence among the six faba bean varieties, particularly for NS and weight traits that were remarkably consistent across water regimes.
Genotype × environment interactions were significant for NTS (3.95%,
P<0.05), NP (8.63%,
P<0.01) and NS (6.85%,
P<0.05), but not for WSS (1.92%, ns). The interaction was highly significant for PY (25.15%,
P<0.001), showing that while certain varieties maintained stable stem and pod production across regimes, yield responses differed markedly depending on the water regime. Blocks within environments contributed minimally (2-8% of total variation) and were only significant for NP (5.82%,
P<0.05) and PY (8.25%,
P<0.05). Error variance ranged from 3.22% (WSS) to 21.70% (PY) and coefficients of variation show acceptable experimental precision (CV 6.44% for NP to 46.80% for PY). H² were moderate to high, from 58.6% for NTS up to 96.70% for WSS, indicating that selection for these traits, especially seed weight and seed number traits, should be effective under varying irrigation (Table 1). High H2 estimates support their use as stable selection criteria, as reported in recent faba bean and legume studies (
Esho and Salih, 2021;
Hiywotu et al., 2023; Boukrouh et al., 2024).
Correlation and path analysis of agronomic traits in faba bean
Results revealed that under IR1, Faba bean yield was slightly correlated with NP, NS and WSS parameters (r = 0.10-0.36;
P<0.05), indicating that optimizing these aspects would influence the overall yield. In contrast, strong correlations were spotted between WSS and both NP and NS (r = 0.7 to 0.75) (
P<0.05). Furthermore, the transition from IR1 to the higher levels (IR2 and IR3) further affected these correlative effects. Regarding these conditions, the correlation between yield and faba bean parameters became stronger (r = 0.46-0.65; P<0.05), while the relationship between the variables NP, NS and WSS was weakened over the IR transition (r = 0.18-0.29;
P<0.05) (Fig 1).
Path analysis revealed marked contrasts in the direct and indirect contributions of yield-related traits under different irrigation regimes (Table 2, Fig 2). Under IR1, WSS had a strong positive total effect on yield (+0.86) despite a large negative direct effect (-1.24), implying substantial positive indirect influence-mainly via NP and NS. NP showed a negative direct effect (-0.88) but a positive total correlation (+0.52), indicating its structural, indirect role. NS (-0.15 direct; -0.37 total) and NTS (-0.39) were essentially non-contributory. Under IR2, WSS again showed a strong positive total correlation (+0.75) despite a negative direct effect (-1.15). NP retained a negative direct effect (-0.82) but a positive total (+0.45), reinforcing its indirect importance. NS (-0.20) remained a weak predictor and NTS continued to be unfavorable. Under IR3, WSS had its most negative direct effect (-1.30) yet kept a considerable total correlation (+0.65) via indirect pathways. NP continued to contribute indirectly (-0.92 direct; +0.40 total). NS (-0.10 direct; -0.30 total) remained of limited relevance, while NTS consistently showed negative effects across regimes.
NP acts as a central mediator: although its direct effect is negative, its consistent, strong indirect contribution across irrigation regimes, especially under moderate irrigation, makes it a valuable criterion for indirect selection. By contrast, NS shows persistently negative direct and total effects and is an unreliable yield predictor, particularly under water stress. NTS is broadly unfavorable, suggesting assimilate diversion to vegetative growth at the expense of reproductive organs.
To understand water influence on yield, correlation analysis was conducted between PY and the main agronomic components under different irrigation regimes. Under IR1, associations between PY and components (NP, NS, WSS) were low (r = 0.10-0.36) though significant (Fig 1), suggesting that under water stress, yield is influenced predominantly by external environmental factors (temperature, soil moisture, drought frequency) rather than morpho-productive traits
(Dong et al., 2019). Liao et al., (2022) reported that soybean yield components are strongly influenced by environmental conditions, sometimes exceeding genetic factors. In contrast,
Semcheddine and Hafsi (2014), found no correlation between cereal yield and grain number under rainfed conditions, while positive relationships appeared under controlled irrigation.
Conversely, under IR2 and IR3, the correlations between PY and NP, NS and WSS become more marked (r = 0.46-0.65) (Fig 1), indicating a better expression of the genetic potential of the genotypes. Increased water availability appears to favor the transfer of resources to reproductive structures, reinforcing the predictive value of morphological traits. These results corroborate those of
Sharifi (2014) and
Osman et al., (2019), who demonstrated that under favorable conditions, yield components are more reliable predictors of productivity. However,
Dogan (2019) reported that irrigation water did not affect 1000-grain weight, highlighting the importance of genetic and environmental context.
One of the most remarkable results concerns WSS, which shows a strongly negative direct effect on yield in all regimes (from -1.24 to -1.30), while retaining a positive and significant overall correlation. This suggests an essentially indirect influence of WSS, notably
via NP and NS. This apparent paradox illustrates the central role of WSS as a pivotal factor in yield construction, despite an unfavorable direct contribution. Similar observations were reported by
Shferaw and Tarekegne (2024) and
Hiywotu et al., (2023), who highlighted the importance of the indirect effects of biomass and harvest index
via complex interactions. NP also emerges as an important mediating variable. Its negative direct effect is counterbalanced by a positive total correlation (up to +0.52), particularly under IR2, suggesting a relay role for the beneficial effects of WSS and NS. In contrast, NS shows consistently negative effects (direct and total), disqualifying it as a priority selection criterion in semi-arid Morocco.
Ouji et al., (2017) demonstrated that under water stress, components like NS or WSS can be strongly affected, compromising their stability as indicators.