Ethylene-nitrogen synergism induces tolerance to copper stress by modulating antioxidant system and nitrogen metabolism and improves photosynthetic capacity in mustard

This study aimed to test the efficiency of ethylene (Eth; 200 µL L^-1 ethephon) in presence or absence of nitrogen (N; 80 mg N kg^-1 soil) in protecting photosynthetic apparatus from copper (Cu; 100 mg Cu kg^-1 soil) stress in mustard (Brassica juncea L.) and to elucidate the physio-biochemical modulation for Eth plus N-induced Cu tolerance. Elevated Cu-accrued reductions in photosynthesis and growth were accompanied by significantly higher Cu accumulation in leaves and oxidative stress with reduced assimilation of N and sulfur (S). Ethylene in coordination with N considerably reduced Cu accumulation, lowered lipid peroxidation, lignin accumulation, and contents of reactive oxygen species (hydrogen peroxide, H₂O₂, and superoxide anion, O₂^•-), and mitigated the negative effect of Cu on N and S assimilation, accumulation of non-protein thiols and phytochelatins, enzymatic, and non-enzymatic antioxidants (activity of ascorbate peroxidase, APX, and glutathione reductase, GR; content of reduced glutathione, GSH, and ascorbate, AsA), cell viability, photosynthesis, and growth. Overall, the effect of ethylene-nitrogen synergism was evident on prominently mitigating Cu stress and protecting photosynthesis. The approach of supplementing ethylene with N may be used as a potential tool to restrain Cu stress, and protect photosynthesis and growth of mustard plants.