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Microorganisms in Toxic Metal-Polluted Soils 339<br />

extremely reducing conditions which can chemically reduce species such<br />

as U(VI) (White and Gadd 1998b).<br />

Bacterial Fe oxidation is ubiquitous in environments with sufficient Fe 2+<br />

and conditions to support bacterial growth such as drainage waters and<br />

tailings piles in mined areas, pyritic and hydric soils (bogs and sediments),<br />

drain pipes and irrigation ditches, and plant rhizospheres. Iron-oxidizers<br />

foundinacidicsoilenvironmentsareacidophilicchemolithotrophs,such<br />

as Thiobacillus ferrooxidans, significant for its role in generating acid mine<br />

drainage (Ewart and Hughes 1991). Fungi also oxidize metals in their environment.<br />

Desert varnish is an oxidized metal layer (patina) a few millimetres<br />

thick found on rocks and in soils of arid and semi-arid regions, and is<br />

believed to be of fungal and bacterial origin.<br />

6<br />

Metalloid Transformations<br />

Main microbial transformations carried out in the soil are reduction and<br />

methylation which can lead to alterations in bioavailability and toxicity.<br />

For selenium, some bacteria can use SeO 2−<br />

4 as a terminal e− acceptor in<br />

dissimilatory reduction and also reduce and incorporate Se into organic<br />

components,e.g.selenoproteins(assimilatoryreduction).Methylationand<br />

subsequent volatilization of methylated selenium derivatives is also a widely<br />

found property of soil bacteria and fungi and may be an important pro-<br />

cess in Se transport from terrestrial to aquatic environments (Dungan and<br />

Frankenberger 1999). Selenate (SeO2− 4 ) and selenite (SeO2− 3 ) can be reduced<br />

to Seo ,withSeO2− 3 reduction appearing more ubiquitous than SeO2− 4 reduction.<br />

However, only SeO2− 4 can support bacterial growth under anaerobic<br />

conditions: SeO2− 4 reduction to Seo is a major sink for Se oxyanions in anoxic<br />

sediments (Oremland et al. 1989; Stolz and Oremland 1999; Oremland and<br />

Stolz 2000). Anaerobic sulphate-reducing bacteria like Desulfovibrio desulfuricans<br />

can reduce selenate/selenite to Seo , but neither oxyanion could be<br />

used for respiratory growth (Tomei et al. 1995). Reduction to Seo can be<br />

considered a detoxification mechanism (Dungan and Frankenberger 1999).<br />

The opposite process of Seo oxidation can occur in soils and sediments<br />

(Dowdle and Oremland 1998; Losi and Frankenberger 1998). It is possible<br />

that Seo oxidation is a similar process to S oxidation, and may be mediated<br />

by heterotrophs and autotrophs (Losi and Frankenberger 1998). In aerobic<br />

soil slurries, Se4+ was the main product with lower amounts of Se6+ being<br />

produced: heterotrophic and autotrophic thiobacilli were believed to be the<br />

active organisms (Dowdle and Oremland 1998).<br />

Methylation of Se is a ubiquitous property of microorganisms and can<br />

occur in soils, sediments and water (Gadd 1993b). Bacteria and fungi

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