Mechanisms of heavy metal cadmium tolerance in plants

Cadmium (Cd) is a strongly phytotoxic heavy metal, which inhibits plant growth and even leads to plant death. The main symptoms of Cd(2+) toxicity to plants are stunting and chlorosis. Plant has developed some functions for Cd(2+) tolerance, which include cell wall binding, chelation with phytochelatins (PCs), compartmentation of Cd(2+) in vacuole, and enrichment in leaf trichomes. However, Cd(2+) tolerance in plant is more likely involved in an integrated network of multiple response processes than several isolated functions cited above. In the network, the processes of sulfur metabolism, antioxidative response, and Cd(2+) transport across plasma and vacuole membrane in plant are closely related with Cd(2+) tolerance in plant. The processes of sulfur uptake, assimilation and sequential sulfur metabolism in plant respond to Cd(2+) stress. The expression of sulfur transporters with varied affinity was changed in different ways under Cd(2+) stress, and the high expression of ATP sulfurylase (APS) and adenosine 5' phosphosulfate reductase (APR), which may help to keep the supply of S(2-) for cysteine (Cys) synthesis. The efficiency of Cys synthesis may function in Cd(2+) detoxification, and the up-regulated expression of Ser acetyltransferase (SAT) and O-acetyl-ser (thiol)-lyase (OASTL) has been found in some Cd(2+) treated plants. Reduced glutathione (GSH) is an important antioxidant and the precursor of PCs, glutamylcysteine synthetase (GCS) and glutathione synthetase (GS) catalyze GSH synthesis from Cys, overexpression of the two enzymes can improve Cd(2+) tolerance in plant. PCs are more important Cd(2+) chelators than metallothioneins (MTs) in plants, and the expression of phytochelatin synthase (PCS) responds to Cd(2+) stress. Plant antioxidative system also contributes to Cd(2+) tolerance. The antioxidative response to Cd(2+)-induced oxidative stress varies in different plants and tissues and is also Cd(2+) concentration dependent, and the Cd hyperaccumulator plants show strong tolerance to oxidative stress. Some genes encoded metal transporters with Cd(2+) substrate specificity at plasma and vacuole membranes, which have been isolated and characterized in recent years. These genes play critical roles in Cd(2+) translocation, allocation, and compartmentation in plants. Despite the great progresses made in the field in recent years, there are still some issues which need further exploration, such as the detail of signal transduction and the responses of gene regulation to Cd(2+), the rhizosphere activation and root adsorption to soil Cd(2+), Cd(2+) trafficking in xylem and phloem, Cd(2+) translocation to fruit and seed, and the possible presence of a high-affinity Cd(2+) transporter in Cd hyperaccumulators.