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Background and aims Sedum alfredii is a recently identified cadmium (Cd) hyperaccumulator of increasing interest for its potential use in phytoextraction. In this study, we examined short-term responses of roots of the Cd hyperaccumulating ecotype (HE) S. alfredii to Cd exposure with comparison with its non-hyperaccumulating ecotype (NHE).



The toxic trace pollutant cadmium (Cd) enters the environment mainly from industrial processes and phosphate fertilizers (Wagner, 1993), and is damaging to plant, animal and human health (Bertina and Averbeck, 2006). In plants, a few micromoles of Cd in the root environment is toxic resulting in root tip damage, reduced photosynthesis, induced antioxidant responses in all plant organs, and growth inhibition (Das et al., 1998). Cadmium is assumed to cause the direct or indirect formation of reactive oxygen species (ROS) in plants (Rodr¨ªguez-Serrano et al., 2006), and to interfere with the redox status of cells (Sandalio et al., 2001; Sch¨¹tzend¨¹bel et al., 2001; Bertina and Averbeck, 2006). In most plants, visible effects of exposure to high Cd doses are growth inhibition and leaf chlorosis (Das et al., 1998).
In some plant species including Thlaspi caerulescens (Boominathan and Doran, 2003) and Brassica juncea (Pilon-Smits et al., 2000), Cd can accumulate to substantial concentrations without any toxic effects. The tolerance of Cd by these species involves a complex network of homeostatic mechanisms that control the uptake, accumulation, trafficking, and detoxification of metals (Das et al., 1998; Clemens, 2001). Once uptake and translocation of Cd have occurred, a number of mechanisms are involved that regulate the concentrations of free Cd ions among different plant organelles and hence minimize the damage to metabolism from exposure to excess Cd (Clemens, 2001; Hall, 2002). Compartmentations to vacuoles and cell walls (Bidwell et al., 2004; Krämer et al., 2000), and chelation with agents (Krämer et al., 1996; Salt et al., 1999; Ueno et al., 2005) have been suggested to play a dominant role in metal detoxification by hyperaccumulators (Clemens, 2001). Metal detoxification, however, is frequently incomplete and mechanisms that protect metabolism from metal-induced oxidative damage are necessarily involved (Boominathan and Doran, 2002; 2003; Freeman et al., 2004). The non-protein thiol (NPT), glutathione (GSH) has been suggested to play an important role in detoxification of heavy metals in plants (Han et al., 2008). GSH-mediated Ni tolerance has been previously observed in Thlaspi hyperaccumulators (Freenman et al., 2004), and Cd tolerance/accumulation was reported to be associated with GSH biosynthesis in the accumulator plant Brassica juncea (Zhu et al., 1999a; 1999b; Pilon-Smits et al., 2000).
Sedum alfredii is a recently identified Cd hyperaccumulators (Yang et al., 2004), of increasing interest (Li et al., 2007) for its potential use in phytoextraction. Plants of the hyperaccumulating ecotype (HE) of S. alfredii grow natrually in a Pb/Zn area, where Cd concentration is up to 400 mg kg-1 soil. Yang et al. (2004) reported that this ecotype of S. alfredii grew healthy at Cd levels of up to 200 µM hydroponicly, whereas its contrasting ecotype (the nonhyperaccumulating ecotype, NHE) can not survive at 50 µM (Xiong et al., 2005). Like other hyperaccumulators, Sedum alfredii has an enhanced ability to translocate most of absorbed Cd from roots to shoots (Lu et al., 2008), to protect the roots from toxicity of excess metals. However, roots themselves must also possess an extraordinary ability to tolerate Cd, as in most environmental conditions the toxic metal enters the roots first. Roots of the Cd hyperaccumulator, Thlaspi caerulescens (Nedelkoska and Doran, 2000), have been suggested to possess mechanisms for enhanced metal tolerance independent of the shoots, and antioxidative metabolism might play an important role in countering the toxic effects of Cd in hairy roots of T. caerulescens (Boominathan and Doran, 2003). Previous studies on the hyperaccumulator S. alfredii indicated that there was no relationship between antioxidative defense mechanisms namely superoxide dismutase, catalase, guaiacol peroxidase, ascorbate peroxidase and glutathione reductase activities as well as ascorbic acid contents, and Cd tolerance in both roots and leaves of S. alfredii (Jin et al., 2008a). The studies by Jin et al., (2008a), however, did not provide a clear explanation for the apparent ability of roots to tolerate high cellular Cd levels. Roots tip, which comprises meristem, elongation and differentiation cells within a short distance (Scheres et al., 2002), is a very active zone of metabolism, and extremely sensitive to biotic or abiotic stress. Therefore, a precise determination for antioxidative capacity of the root tips is necessary to better understand the tolerance mechanism of HE S. alfredii in the early response to Cd.

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