Uptake, impact, adaptive mechanisms, and phytoremediation of heavy metals by plants: Role of transporters in heavy metal sequestration.

Heavy metals (HMs) pose severe threats to both the environment and its inhabitants, leading to reduced crop productivity and hazardous impacts on human and animal health. Metallurgical activities in peri-urban areas are major contributors to the terrestrial deposition of various HMs. Upon entering plant the cells, HMs disrupt structural and physiological processes, inducing stress responses and triggering metabolic pathways for stress adaptations. The plants have evolved specialized transport systems to regulate the uptake, transport, and cellular concentrations of these metals. HMs often exploit transporters of essential nutrients, such as phosphate, hexose, and sulfate to gain entry into plant cells. Key players include zinc receptor transporter (ZRT1) and iron receptor transporter (IRT1), both part of the ZIP (Zinc Iron Permease) family, as well as heavy metal-associated ATPases (HMAs) and ATP binding cassette transporter C (ABCC-type transporters). Hyperaccumulating plants thrive in harsh environments with elevated concentrations of toxic ions, such as sodium, chloride, and heavy metals including arsenic (As), mercury (Hg), cadmium (Cd), lead (Pb), silicon (Si), boron (B), antimony (Sb), germanium (Ge), and tellurium (Te), by compartmentalizing these ions into vacuoles. The accumulation of heavy metals or metalloids like cadmium (Cd), lead (Pb), arsenic (As), chromium (Cr), nickel (Ni), manganese (Mn), zinc (Zn), thallium (Tl), cobalt (Co), cupper (Cu), and selenium (Se) has been extensively reported in various hyperaccumulating plant species. The halophytes, known for their inherent salinity tolerance, exhibit superior resilience to HM stress due to overlapping mechanisms of ion compartmentatlization and detoxification. This review provides an in-depth analysis on the effects of heavy metals on the metabolic processes, growth, and development of plants, emphasizing heavy tolerance mechanisms with a particular focus on halophytes. The role of HM transporters in metal sequestration and detoxification is discussed, along with the potential of hyperaccumulating halophytes for phytoremediation of HM-contaminated soils.