Combined Effect of Salicylic Acid and Proline Mitigates Drought Stress in Rice (Oryza sativa L.) through the Modulation of Physiological Attributes and Antioxidant Enzymes

Salicylic acid (SA) and proline exhibit protective effects against a wide range of stresses. However, the combined impact of SA and proline on rice under drought stress is still unknown. Therefore, we investigated the protective roles of SA and/or proline in conferring drought tolerance in rice. There were eight treatments comprising the control (T1; 95–100% FC), 1.5 mM SA (T2), 2 mM proline (T3), 0.75 mM SA + 1 mM proline (T4), 45–50% FC (T5, drought stress), T5 + 1.5 mM SA (T6), T5 + 2 mM proline (T7), and T5 + 0.75 mM SA + 1 mM proline (T8), and two rice varieties: BRRI dhan66 and BRRI dhan75. Drought stress significantly decreased the plant growth, biomass, yield attributes, photosynthetic rate (Pn), stomatal conductance (gs), transpiration rate (Tr), photosynthetic pigments (chlorophyll and carotenoids content), relative water content (RWC), membrane stability index (MSI), soluble sugar and starch content, and uptake of N, P and K(+) in roots and shoots. Drought-induced oxidative stress in the form of increased hydrogen peroxide (H(2)O(2)) production and lipid peroxidation (MDA) was observed. The combined application of SA (0.75 mM) + proline (1 mM) was found to be more effective than the single application of either for drought stress mitigation in rice. A combined dose of SA + proline alleviated oxidative stress through boosting antioxidant enzymatic activity in contrast to their separate application. The application of SA + proline also enhanced proline, soluble sugar and starch content, which resulted in the amelioration of osmotic stress. Consequently, the combined application of SA and proline significantly increased the gas exchange characteristics, photosynthetic pigments, RWC, MSI, nutrient uptake, plant growth, biomass and yield of rice. Therefore, the combined application of SA and proline alleviated the detrimental impacts of drought stress more pronouncedly than their separate application did by increasing osmoprotectants, improving nutrient transport, up-regulating antioxidant enzyme activity and inhibiting oxidative stress.