Physiological and biochemical bases of spermidine-induced alleviation of cadmium and lead combined stress in rice.

Cadmium (Cd) and lead (Pb) pollution is a major environmental issue affecting plant production. Spermidine (Spd) is involved in plant response to abiotic stress. However, the role and associated mechanism of Spd under Cd + Pb combined stress are poorly understood. The potential protective role of Spd at different concentration on rice (Oryza sativa L.) seedlings exposed to Cd + Pb treatment was investigated by a hydroponic experiment in this study. The results showed that exogenous Spd enhanced the tolerance of rice seedlings to Cd + Pb stress, resulted in an increase in plant height, root length, fresh weight and dry weight of roots and shoots. Further, application of Spd decreased the contents of hydrogen peroxide, superoxide anion, malondialdehyde, and the accumulation of Cd and Pb, and increased the contents of mineral nutrient, carotenoids, chlorophyll, proline, soluble sugar, soluble protein, total phenol, flavonoid, anthocyanin, and antioxidant enzymes activities in roots and shoots of rice seedlings under Cd + Pb stress. Particularly, 0.5 mmol L Spd was the most effective to alleviate the adverse impacts on growth and physiological metabolism of rice seedlings under Cd + Pb stress. Principal component analysis and heat map clustering established correlations between physio-biochemical parameters and further revealed Spd alleviated Cd + Pb damage in rice seedling was associated with inhibition of accumulation and translocation of Cd and Pb, increasing the contents of photosynthetic pigments and mineral nutrient and stimulation of antioxidative response and osmotic adjustment. Overall, our findings provide an important prospect for use of Spd in modulating Cd + Pb tolerance in rice plants. Spd could help to alleviate Cd + Pb damage through inhibition of accumulation and translocation of Cd and Pb and stimulation of oxidant-defense system and osmotic adjustment.