Significance of vacuolar proton pumps and metal/H antiporters in plant heavy metal tolerance.

Soil and water are among the most valuable resources on earth. Unfortunately, their contamination with heavy metals has become a global problem. Heavy metals are not biodegradable and cannot be chemically degraded; therefore, they tend to accumulate in soils or to be transported by streaming water and contaminate both surface and groundwater. Cadmium (Cd) has no known biological function but is one of the most toxic metals. It represents a serious environmental concern since its accumulation in soils is associated with health risks to plants, animals and humans. On the other hand, copper (Cu) and zinc (Zn) are heavy metals that are indispensable to plants but become toxic when their concentration in soils exceeds a certain optimal level. Plants have evolved many mechanisms to cope with heavy metal toxicity; vacuolar sequestration is one of them. Vacuolar sequestration can be achieved through either phytochelatin-dependent or phytochelatin-independent pathways. Most of the transgenic plants meant for phytoremediation described in the literature result from the manipulation of genes involved in the phytochelatin-dependent pathway. However, recent evidence has emerged to support the importance of the phytochelatin-independent pathway in heavy metal sequestration into the vacuole, with metal/H antiporters and proton pumps playing an important role. In this review, the importance of vacuolar proton pumps and metal/H antiporters transporting Cd, Cu, and Zn is discussed. In addition, the recent advances in the production of transgenic plants with potential application in phytoremediation and food safety through the manipulation of genes encoding V-PPase proton pumps is described.