Water deficit aggravated the inhibition of photosynthetic performance of maize under mercury stress but is alleviated by brassinosteroids

Mercury (Hg) significantly inhibits maize (Zea mays L.) production, which could be aggravated by water deficit (WD) due to climate change. However, there is no report on the maize in response to combined their stresses. This work was conducted for assessing the response and adaptive mechanism of maize to combined Hg and WD stress using two maize cultivars, Xianyu (XY) 335 and Yudan (YD) 132. The analysis was based on plant growth, physiological function, and transcriptomic data. Compared with the single Hg stress, Hg accumulation in whole plant and translocation factor (TF) under Hg+WD were increased by 64.51 % (1.44 mg kg-1) and 260.00 %, respectively, for XY 335; and 50.32 % (0.62 mg kg-1) and 220.02 %, respectively, for YD 132. Combined Hg and WD stress further increased the reactive oxygen species accumulation, aggravated the damage of the thylakoid membrane, and decreased chlorophyll content compared with single stress. For example, Chl a and Chl b contents of XY 335 were significantly decreased by 48.67 % and 28.08 %, respectively at 48 h after Hg+WD treatment compared with Hg stress. Furthermore, transcriptome analysis revealed that most of down-regulated genes were enriched in photosynthetic-antenna proteins, photosynthesis, chlorophyll and porphyrin metabolism pathways (PsbS1, PSBQ1 and FDX1 etc.) under combined stress, reducing light energy capture and electron transport. However, most genes related to the brassinosteroids (BRs) signaling pathway were up-regulated under Hg+WD stress. Correspondingly, exogenous BRs significantly enhanced the maize tolerance to stress by decreasing Hg accumulation and TF, and raising activities of antioxidant enzyme, the content of chlorophyll and photosynthetic performance. The PI, Fv/Fm and Fv/Fo of Hg+WD+BR treatment were increased by 29.88 %, 32.06 %, and 14.56 %, respectively, for XY 335 compared to Hg+WD. Overall, combined Hg and WD stress decreased photosynthetic efficiency by adversely affecting light absorption and electron transport, especially in stress-sensitive variety, but BRs could alleviate the inhibition of photosynthesis, providing a novel strategy for enhancing crop Hg and WD tolerance and food safety.