TILLAGE & CULTIVATION·PHYSIOLOGY & BIOCHEMISTRY·AGRICULTURE INFORMATION TECHNOLOGY
WANG Xue-hua, DAI Li
A certain degree of cadmium stress has seriously influenced crop growth, development, yield and quality of farm produce. In this review, the authors comprehensively summarized the damages of Cd to crops and a human being, and the characteristics of the Cd absorption, transport and accumulation as well as their dependent main regulatory genes and their functions. The resistant and tolerant mechanisms of crops to the Cd toxicity were simply summarized, while the physiology and biochemical mechanism of the root cadmium immobilization effect which was one of them were emphatically introduced. The Cd got into the plant via the root absorption mostly, and in the root, the Cd2+ entered the “free space” firstly, which is constituted by the cell space, the cell wall micropore and the space between cell wall and plasma membrane, afterwards, passed through the plasma membrane into the cytoplasm by the way of active or passive absorption, and then were transported into the xylem vessel through symplast or apoplast pathway. Rice and others crops mainly use the following ways to adapt the Cd stress: the retention effect of cell wall, the chelation effect in symplast, the compartmentation effect of vacuole, the accumulation of stress protein and proline, the enhancement of antioxidase system activity and the immobilization effect of the root. As an important way to resist and tolerate the cadmium toxicity, the immobilization effect of cadmium in crop root plays a crucial role in regulating the Cd absorption, transport, distribution and accumulation in crop, preventing Cd entering into bioplasts and shoots of plant, and reducing the harm of Cd to the growth and development of the crop and the yield and quality of the farm produce. It mainly contains the retention caused by the low root-to-shoot Cd transport volume, the cell wall and the vacuolar. (i) The retention caused by the low root-to-shoot Cd transport volume. This retention effect is mostly influenced by the xylem Cd loading capacity of the root and the content of the cadmium long distance transport carrier – phytochelatins (PCs); and they were controlled by the ion transporter HMA2 and HMA4 on plasma membrane and the PCs synthetase in cell and their relevant regulatory genes, such as HMA2, HMA4 and PCs1. And they have a negative regulation to the Cd retention in the xylem. (ii) The cell wall immobilization effect. This effect happens in the apoplast (including the cell wall and intercellular layer) of the root cell, and is mainly related to the component and structure of the apoplast, and in which the pectin has played a key role, moreover, the hemicellulose also has some effect in it. On the basis of the different action way in the pectin and hemicellulose Cd retention, the cell wall immobilization effect can be divided into physical immobilization (PIB) and chemical immobilization (CIB). The PIB is mostly related to the pectin content and the pectin methyl esterase (PME) activity. While, the CIB is the result of electrostatic binding effect between the Cd2+ and carboxyl group or other groups with negative charge on the pectin, hemicellulose and other components of the cell wall. And they were controlled by the genes like PME14 and XCD1. (iii) The vacuole immobilization effect. This effect is closely related to the PCs in the cytoplasm and the vacuole, as well as the transport protein on the tonoplast; and its retention ability to Cd is restricted by the vacuolar sequestration capacity (VSC). In the vacuole Cd immobilization, the PCs with different molecular weight play an important part, they participated in the chelation of Cd in the cytoplasm, the Cd transformation between cytoplasm and vacuole, and the Cd sedimentation in the vacuole finally. At the same time, the transport proteins on the tonoplast are in charge of transferring the low molecular weigh PC-Cd compound from cytoplasm solution into the vacuole through active transport, making the Cd insulated. In crop roots, the successively combined action of the three immobilization effects has reduced the cadmium transfer amount from the root to shoot, then eased the pernicious effects of the Cd to the shoot, and decreased the content in the crop harvesting organs like grain. However, due to the finiteness of the percentage of PC-Cd in the general Cd number, the total charge number of the cell wall and the VSC of vacuole, making the intensity and effectiveness of the root-Cd-retention effects are limited to a certain degree.