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The abilities to mobilize and/or sequester excess ions within and outside the plant cell are important components of salt-tolerance mechanisms. Mobilization and sequestration of Na +  involves three transport systems facilitated by the plasma membrane H + /Na +  antiporter (SOS1), vacuolar H + /Na +  antiporter (NHX1), and Na + /K +  transporter in vascular tissues (HKT1). Many of these mechanisms are conserved across the plant kingdom. While  Gossypium hirsutum  (upland cotton) is significantly more salt-tolerant relative to other crops, the critical factors contributing to the phenotypic variation hidden across the germplasm have not been fully unraveled. In this study, the spatio-temporal patterns of Na +  accumulation along with other physiological and biochemical interactions were investigated at different severities of salinity across a meaningful genetic diversity panel across cultivated upland  Gossypium.  The aim was to define the importance of holistic or integrated effects relative to the direct effects of Na +  homeostasis mechanisms mediated by  GhHKT1, GhSOS1 , and  GhNHX1.  Multi-dimensional physio-morphometric attributes were investigated in a systems-level context using univariate and multivariate statistics,  randomForest , and path analysis. Results showed that mobilized or sequestered Na +  contributes significantly to the baseline tolerance mechanisms. However, the observed variance in overall tolerance potential across a meaningful diversity panel were more significantly attributed to antioxidant capacity, maintenance of stomatal conductance, chlorophyll content, and divalent cation (Mg 2+ ) contents other than Ca 2+  through a complex interaction with Na +  homeostasis. The multi-tier macro-physiological, biochemical and molecular data generated in this study, and the networks of interactions uncovered strongly suggest that a complex physiological and biochemical synergy beyond the first-line-of defense (Na +  sequestration and mobilization) accounts for the total phenotypic variance across the primary germplasm of  Gossypium hirsutum . These findings are consistent with the recently proposed Omnigenic Theory for quantitative traits and should contribute to a modern look at phenotypic selection for salt tolerance in cotton breeding.

frontiers in plant science

Networks of Physiological Adjustments and Defenses, and Their Synergy With Sodium (Na+) Homeostasis Explain the Hidden Variation for Salinity Tolerance Across the Cultivated Gossypium hirsutum Germplasm