MdbHLH93 confers drought tolerance by activating MdTyDC expression and promoting dopamine biosynthesis

Dopamine and its biosynthesis-limiting enzyme tyrosine decarboxylase (TyDC) play a vital part in mediating plant growth and the response to drought stress. However, the underlying molecular mechanism remains poorly understood. Here, drought stress markedly induced the expression of Malus domestica bHLH93 (MdbHLH93), the apple basic helix-loop-helix transcription factor. Moreover, MdbHLH93 directly bound to the Malus domestica TyDC (MdTyDC) promoter and positively regulated its expression, which resulted in dopamine synthesis and enhanced drought tolerance. Furthermore, the additive effect of overexpressing MdbHLH93 and MdTyDC simultaneously promoted dopamine synthesis and drought tolerance in apples, while the interference of MdbHLH93 inhibited this effect, indicating that MdTyDC-regulated dopamine synthesis and drought tolerance were positively regulated by MdbHLH93. Taken together, these findings suggest the positive regulation of dopamine accumulation by MdbHLH93 through its transcriptional regulation of MdTyDC and show that increased dopamine content confers drought tolerance in apples.