Role of phytohormones in heavy metal tolerance in plants: A review

Heavy metal (HM)-mediated toxic effects on plants have attained considerable attention worldwide as they directly threaten the food supply chain. Although various measures have been taken to mitigate the adverse effects of heavy metal stress in plants, significant research gaps must be proactively addressed. Fascinatingly, the exogenous application of phytohormones has recently attained substantial interest in regulating the negative effects of HM stress. Phytohormones are signaling transductional molecules that mitigate HM toxicity in plants and support their growth and development. Both exogenous treatments and manipulation of the endogenous status of phytohormones through regulating their signaling/biosynthesis-related genes are effective strategies for protecting plants against HM-induced toxic effects. However, to achieve maximum benefits from phytohormone-mediated subcellular mechanisms to mitigate HM toxicity, it is necessary to gain in-depth understanding on their potential pathways. The current review focuses primarily on the major mechanistic phytohormonal-mediated approaches involved in alleviating the toxic effects of HMs in plants. Moreover, the potential roles of major phytohormones in triggering protein molecules, signaling transductions, and gene expressions to avoid, tolerate, or alleviate HM toxicity in plants are also discussed. This information provides systematic understanding on the mechanisms of phytohormones in modulating heavy metal tolerance in plants and could help to guide the development of strategies to improve plant tolerance against HM toxicity.