Alleviation of Adverse Effects of Drought Stress on Growth and Nitrogen Metabolism in Mungbean (Vigna radiata) by Sulphur and Nitric Oxide Involves Up-Regulation of Antioxidant and Osmolyte Metabolism and Gene Expression

The influence of drought induced by polyethylene glycol (PEG) and the alleviatory effect of nitric oxide (50 µM) and sulphur (S, 1 mM K(2)SO(4)) were studied in Vigna radiata. Drought stress reduced plant height, dry weight, total chlorophylls, carotenoids and the content of nitrogen, phosphorous, potassium and sulphur. The foliar applications of NO and sulphur each individually alleviated the decline, with a greater alleviation observed in seedlings treated with both NO and sulphur. The reduction in intermediates of chlorophyll synthesis pathways and photosynthesis were alleviated by NO and sulphur. Oxidative stress was evident through the increased hydrogen peroxide, superoxide and activity of lipoxygenase and protease which were significantly assuaged by NO, sulphur and NO + sulphur treatments. A reduction in the activity of nitrate reductase, glutamine synthetase and glutamate synthase was mitigated due to the application of NO and the supplementation of sulphur. The endogenous concentration of NO and hydrogen sulphide (HS) was increased due to PEG; however, the PEG-induced increase in NO and HS was lowered due to NO and sulphur. Furthermore, NO and sulphur treatments to PEG-stressed seedlings further enhanced the functioning of the antioxidant system, osmolytes and secondary metabolite accumulation. Activities of γ-glutamyl kinase and phenylalanine ammonia lyase were up-regulated due to NO and S treatments. The treatment of NO and S regulated the expression of the Cu/ZnSOD, POD, CAT, RLP, HSP70 and LEA genes significantly under normal and drought stress. The present study advocates for the beneficial use of NO and sulphur in the mitigation of drought-induced alterations in the metabolism of Vigna radiata.