“…In contrast to metal excluders whose strategy is to control the uptake of metals into the root and prevent metal translocation to aerial organs, hyperaccumulators accumulate metals in the shoot to levels toxic to most other plants (Baker, 1981;Baker and Brooks, 1989;Baker et al, 2000;Pollard et al, 2002;Krämer, 2010;Rascio and Navari-Izzo, 2011). This is remarkable as the photosynthetic apparatus is one of the major targets of metal phytotoxicity, typically resulting in severe symptoms such as chlorosis and necrosis, wilting, abnormal development and reduced growth (Pandey and Sharma, 2002;Rahman et al, 2005;Marschner and Marschner, 2012). These toxic effects are a product of numerous harmful interactions at the cellular level (Haydon and Cobbett, 2007), including nonspecific binding of metals to enzyme functional groups and displacement of other metals from their binding sites, generation of reactive oxygen species by redox-active metals that can lead to disruption of the electrontransport chain (Qadir et al, 2004), lipid peroxidation and subsequent impairment of membrane integrity (Pandolfini et al, 1992;Ros et al, 1992;Gonnelli et al, 2001;Haydon and Cobbett, 2007;Krämer, 2010;Hanikenne and Nouet, 2011).…”