Manganese enhances cadmium tolerance in barley through mediating chloroplast integrity, antioxidant system, and HvNRAMP expression

Cadmium (Cd) is a toxic heavy metal that poses risks to crop production and food safety worldwide. This study evaluated whether manganese (Mn) addition could mitigate Cd toxicity and reduce Cd accumulation in barley seedlings. Hydroponically grown seedlings of Cd-tolerant (WSBZ) and Cd-sensitive (Dong17) barley cultivars were treated with 0.1 μM and 1 μM Cd as well as 0.2 mM Mn alone and in a combination with 0.1 or 1.0 μM Cd for 21 days. Cd exposure caused the dramatic alteration of growth and physiological parameters by disrupting chloroplast, and increased Cd accumulation in both genotypes. However, Mn addition markedly alleviated the negative impacts of all examined parameters caused by Cd stress. Cd addition enhanced expression of anti-oxidative enzyme related genes, including HvSOD, HvCAT, HvAPX, HvPOD in the two barley genotypes exposed to Cd stress. The expression analysis showed nearly all HvNRAMPs genes are dramatically up regulated by both Mn and Cd, with WSBZ having higher expression than Dong 17. Notably, HvNRAMP1 showed the highest expression due to Mn addition, highlighting its crucial role in Mn uptake and transportation in barley. Moreover, Cd stress and Mn addition increased and suppressed the expression of HvYSL5, HvHMA2 and HvHMA3, respectively. Conversely, the expression of HvYSL2, HvIRT1 and HvMTP8 was upregulated by both Mn and Cd treatments, with a further increase observed in the combined Cd and Mn treatments. It may be concluded that sufficient Mn supply is quite important for reducing Cd uptake and accumulation in plants.