b Selenium species, particularly the oxyanions selenite (SeO 3 2؊ ) and selenate (SeO 4 2؊ ), are significant pollutants in the environment that leach from rocks and are released by anthropogenic activities. Selenium is also an essential micronutrient for organisms across the tree of life, including microorganisms and human beings, particularly because of its presence in the 21st genetically encoded amino acid, selenocysteine. Environmental microorganisms are known to be capable of a range of transformations of selenium species, including reduction, methylation, oxidation, and demethylation. Assimilatory reduction of selenium species is necessary for the synthesis of selenoproteins. Dissimilatory reduction of selenate is known to support the anaerobic respiration of a number of microorganisms, and the dissimilatory reduction of soluble selenate and selenite to nanoparticulate elemental selenium greatly reduces the toxicity and bioavailability of selenium and has a major role in bioremediation and potentially in the production of selenium nanospheres for technological applications. Also, microbial methylation after reduction of Se oxyanions is another potentially effective detoxification process if limitations with low reaction rates and capture of the volatile methylated selenium species can be overcome. This review discusses microbial transformations of different forms of Se in an environmental context, with special emphasis on bioremediation of Se pollution.
Since the discovery in 1954 by Pinsent that the oxidation of formate by cell suspensions of Escherichia coli requires growth medium containing molybdate and selenite, there has been a growing interest in the biochemical role of selenium in microorganisms (1). Se is an essential component of selenoamino acids, such as selenomethionine and selenocysteine (the 21st proteinogenic amino acid), that occur in certain types of prokaryotic enzymes. Indeed, the requirement for selenite in E. coli growing on formate is linked to the fact that formate dehydrogenase contains selenocysteine. Other prokaryotic enzymes that contain selenocysteine include glycine reductase in several clostridia, formate dehydrogenases in diverse prokaryotes, including Salmonella, Clostridium, and Methanococcus, as well as hydrogenases in Methanococcus and other anaerobes. In addition, other bacterial Se-dependent enzymes, in which the selenium is part of the active site molybdenum-containing cofactor, include nicotinic acid dehydrogenase and xanthine dehydrogenase, which is present in certain clostridial species (2-4).Reactions that are involved in the cycling of Se in soil, including those influenced by microbes, are diagrammatically summarized in Fig. 1 2Ϫ and was spatially separated from sulfate reduction in the environment despite the presence of substantial concentrations of sulfate where it occurred. Thus, it can be concluded that Se and S have different reductive biogeochemical cycles and appear to involve distinct populations of microorganisms.With respect to the remediation of sel...