400 hyperaccumulator species have been identified, according to the analysis of field-collected specimens. Metal hyperaccumulators are interesting model organisms to study for the development of a phytoremediation technology, the use of plants to remove pollutant metals from soils. However, little is known about the molecular, biochemical, and physiological processes that result in the hyperaccumulator phenotype. We investigated the role of Ni tolerance and transport in Ni hyperaccumulation by Thlaspigoesingense, using plant biomass production, evapotranspiration, and protoplast viability assays, and by following short-and long-tem uptake of Ni into roots and shoots. As long as both species (T. goesingense and Thlaspi arvense) were unaffected by Ni toxicity, the rates of Ni translocation from roots to shoots were the same in both the hyper-and nonaccumulator species. Our data suggest that Ni tolerance is sufficient to explain the Ni hyperaccumulator phenotype observed in hydroponically cultured T. goesingense when compared with the Ni-sensitive nonhyperaccumulator T. arvense.
~~~~ ~~~~~~~~~~~~~~~~~~The Ni requirement of plants is generally very low, 1.7 nmol g-' Ni or less in tissue dry biomass (Brown et al., 1988;Dalton et al., 1988). Symptoms of Ni toxicity can be observed between 0.19 and 0.85 pmol g-' Ni in plant dry biomass. These symptoms include the inhibition of root elongation and interveinal chlorosis, the latter possibly a consequence of the interference of Ni with chlorophyll formation (Woolhouse, 1983;Gabbrielli et al., 1990;Brune and Dietz, 1995;Marschner, 1995). The majority of plants that occur on metalliferous soils are known to exclude toxic metals from their shoots (Baker and Walker, 1990). A number of species, however, have developed an unusual adaptation to metal-rich soils. Instead of excluding toxic metals, so-called hyperaccumulators accumulate metals such as Ni, Zn, or Co in their aboveground biomass. This trait is