A comparative inventory of metal transporters in the green alga Chlamydomonas reinhardtii and the red alga Cyanidioschizon merolae.

As in all organisms, metal cations are crucial for nutrition in plants. Severalmetals, such as copper, iron, zinc, and manganese, act as important cofactors for many enzymes and are essential for bothmitochondrial and chloroplast functions. However, when supplied in excess, these essential cations can become toxic, like heavy metals with no generally established function, such as cadmium, lead, or mercury. Tomaintain micronutrient metal homeostasis and to cope with the deleterious effects of nonessential heavy metals, plants have developed a complex network of metal uptake, chelation, trafficking, and storage processes. Metal transporters are required to maintain metal homeostasis and thus constitute important components of this network (Clemens, 2001; Hall and Williams, 2003). In recent years, a number of membrane transport protein families have been implicated in metal homeostasis in plants. These include the cation diffusion facilitators (CDF), the Zrt-, Irt-like proteins (ZIP), the cation exchangers (CAX), the copper transporters (COPT), the heavy-metal P-type ATPases (HMA), the natural resistance-associated macrophage proteins (NRAMP), and the ATP-binding cassette (ABC) transporters (Williams et al., 2000;Maser et al., 2001; Cobbett et al., 2003;Hall andWilliams, 2003). These transporters are encoded bymultigene families. For example, 15ZIP genes, 12 metal tolerance protein (MTP) genes, and 8 HMA genes are present in the Arabidopsis (Arabidopsis thaliana) genome (Cobbett et al., 2003; Delhaize et al., 2003). However, the transport specificities, patterns of expression, or subcellular localizations of metal transport proteins are still largely unknown. To further understand plant metal homeostasis, it will be necessary to elucidate the contribution of each of these transporters to the uptake, trafficking, and storage of essential metals, as well as to the detoxification of toxic heavy metals. In this article, we present an overview of our current knowledge of the metal transport function in metal homeostasis and tolerance in eukaryotes, with a special emphasis on plants. We also provide a timely inventory of putative metal transporters in two unicellular algal models, the green alga Chlamydomonas reinhardtii and the red alga Cyanidioschizon merolae. These new data should facilitate functional genomics and molecular analysis of metal homeostasis and tolerance in photosynthetic organisms. Moreover, the comparison of metal transporters from species belonging to the red and green algae with those of the land plant Arabidopsis, as well as with their human and yeast homologs, allows some light to be shed on the molecular evolution of metal homeostasis and tolerance systems.