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Abstract <p>Salinity is a major abiotic constraint affecting agricultural productivity, yet limited research has integrated physiological, genetic, and environmental factors to explain resilience in woody Mediterranean species. This study provides the first integrative assessment of <i>Ceratonia siliqua</i> L. (carob) co-cultivated with the halophyte <i>Spergularia salina</i> for salt stress mitigation, combining multivariate analysis (MANOVA, PCA), Partial Least Squares Structural Equation Modeling (PLS-SEM), and AI-assisted hierarchical Bayesian inference. Six Moroccan carob ecotypes (Agadir, Khemissat, Ouazzane, Safi, Aït Attab, Berkane) were evaluated under four NaCl concentrations (0, 85, 171, 257 mM) in monoculture and in co-culture with <i>S.&#xa0;salina</i>. Parameters measured included biomass, aerial length, root length, relative water content (RWC), total chlorophyll, proline, total soluble sugars, and soil electrical conductivity (EC). Salinity significantly increased EC (up to +641.9% in Ouazzane), while co-culture reduced EC under moderate to high salinity (C2H, C3H). Biomass decreased by up to 83% at 257 mM NaCl, with Khemissat most affected, but co-culture improved biomass by up to 80.4% (Berkane). Root length declined by up to 50% under stress, yet co-culture increased it by 67.7% (Agadir). RWC fell by 25–48%, chlorophyll decreased by 34–50%, while proline and total sugars increased markedly, with Ouazzane sugars rising by 226%. MANOVA revealed highly significant effects of both salt stress and ecotype (<i>P</i> &lt; 2.2e-16) on all parameters. PLS-SEM identified root length as the main driver of biomass under salinity (β = 0.629, <i>P</i> &lt; 0.05). Bayesian modeling separated genetic (G[i]) from environmental (E[i]) effects, highlighting Ouazzane as the most tolerant ecotype, Safi and Aït Attab as moderately tolerant, and Khemissat as the most sensitive. This study addresses the gap in integrated genetic–environmental modeling for perennial crops under salinity and demonstrates the potential of combining varietal selection with halophyte co-culture as an agroecological strategy to enhance resilience and support sustainable management of saline soils.</p>

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Genetic and Environmental Determinants of Carob Plants (Ceratonia siliqua L.) Resilience to Salt Stress: Integrative Multivariate, PLS-SEM and Bayesian Modelling

  • 期刊论文

  • 发表日期：

    2026年5月7日

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  • Y. Mouniane,
  • M. El Bakkali,
  • Amol D. Vibhute,
  • I. El-Khadir,
  • A. Chriqui,
  • D. Hmouni

摘要

Abstract

Salinity is a major abiotic constraint affecting agricultural productivity, yet limited research has integrated physiological, genetic, and environmental factors to explain resilience in woody Mediterranean species. This study provides the first integrative assessment of Ceratonia siliqua L. (carob) co-cultivated with the halophyte Spergularia salina for salt stress mitigation, combining multivariate analysis (MANOVA, PCA), Partial Least Squares Structural Equation Modeling (PLS-SEM), and AI-assisted hierarchical Bayesian inference. Six Moroccan carob ecotypes (Agadir, Khemissat, Ouazzane, Safi, Aït Attab, Berkane) were evaluated under four NaCl concentrations (0, 85, 171, 257 mM) in monoculture and in co-culture with S. salina. Parameters measured included biomass, aerial length, root length, relative water content (RWC), total chlorophyll, proline, total soluble sugars, and soil electrical conductivity (EC). Salinity significantly increased EC (up to +641.9% in Ouazzane), while co-culture reduced EC under moderate to high salinity (C2H, C3H). Biomass decreased by up to 83% at 257 mM NaCl, with Khemissat most affected, but co-culture improved biomass by up to 80.4% (Berkane). Root length declined by up to 50% under stress, yet co-culture increased it by 67.7% (Agadir). RWC fell by 25–48%, chlorophyll decreased by 34–50%, while proline and total sugars increased markedly, with Ouazzane sugars rising by 226%. MANOVA revealed highly significant effects of both salt stress and ecotype (P < 2.2e-16) on all parameters. PLS-SEM identified root length as the main driver of biomass under salinity (β = 0.629, P < 0.05). Bayesian modeling separated genetic (G[i]) from environmental (E[i]) effects, highlighting Ouazzane as the most tolerant ecotype, Safi and Aït Attab as moderately tolerant, and Khemissat as the most sensitive. This study addresses the gap in integrated genetic–environmental modeling for perennial crops under salinity and demonstrates the potential of combining varietal selection with halophyte co-culture as an agroecological strategy to enhance resilience and support sustainable management of saline soils.