Aldose reductase expression contributes in sorbitol accumulation and 4-hydroxynon-2-enal detoxification in two foxtail millet (Setaria italica L.) cultivars with different salt stress tolerance

Salt stress is a major environmental factor in arid and semi-arid regions and influences many aspects of plant development. Salinity results in generation of various free radicals that can potentially damage the cellular constituents in plants. Plants were able to effectively reduce the damage caused by these free radicals by a way of enzymatic and non enzymatic defenses for better survival. Enhanced efficacy of antioxidative enzyme systems such as superoxide dismutase, catalase and ascarbate peroxidase was well documented in several plants subjected to salinity stress. Aldose reductase, an important enzyme is also known to detoxify free toxic aldehydes like HNE (4-hydroxynon-2-enal, a hydroxyalkenal) generated during oxidative damage of cellular components. However, the role of aldose reductase to impart tolerance to the plants under salt stress has not been studied in any detail. Therefore, we were interested to study the aldose reductase activity and its expression to gain an insight into the role of aldose reductase in imparting tolerance to foxtail millet cultivars (viz., Cv. Prasad and Lepakshi) subjected to NaCl stress. We observed that subjecting foxtail millets to increasing levels of stress significantly increased aldose reductase activity and in a way that correlated positively with elevated levels of sorbitol, an osmotic solute involved in osmotic balance. This suggests the involvement of aldose reductase in sorbitol biosynthesis in foxtail millet. Additionally, we observed higher levels of 4-hydroxynon-2-enal, a major lipid peroxidation product, in the susceptible than the tolerant cultivar indicating a higher proportion of cellular damage in former than in the latter. This high content of 4-hydroxynon-2-enal in the susceptible cultivar was negatively correlated with its aldose reductase activity, indicating the involvement of aldose reductase in detoxification of 4-hydroxynon-2-enal. 4-hydroxynon-2-enal is also known to be a catalyzed by glutathione-S-transferase. Glutathione-S-transferase activity was found higher in the tolerant foxtail millet than the sensitive cultivar: the tolerant cultivar showed a low 4-hydroxynon-2-enal content compared to the susceptible cultivar, demonstrating a possible mechanism for detoxification of 4-hydroxynon-2-enal by two enzymes, glutathione-S-transferase and aldose reductase in plants under stressful conditions.