In earlier studies, the assimilation of selenate by plants appeared to be limited by its reduction, a step that is thought to be mediated by ATP sulfurylase. Here, the Arabidopsis APS1 gene, encoding a plastidic ATP sulfurylase, was constitutively overexpressed in Indian mustard (Brassica juncea). Compared with that in untransformed plants, the ATP sulfurylase activity was 2- to 2.5-fold higher in shoots and roots of transgenic seedlings, and 1.5- to 2-fold higher in shoots but not roots of selenate-supplied mature ATP-sulfurylase-overexpressing (APS) plants. The APS plants showed increased selenate reduction: x-ray absorption spectroscopy showed that root and shoot tissues of mature APS plants contained mostly organic Se (possibly selenomethionine), whereas wild-type plants accumulated selenate. The APS plants were not able to reduce selenate when shoots were removed immediately before selenate was supplied. In addition, Se accumulation in APS plants was 2- to 3-fold higher in shoots and 1.5-fold higher in roots compared with wild-type plants, and Se tolerance was higher in both seedlings and mature APS plants. These studies show that ATP sulfurylase not only mediates selenate reduction in plants, but is also rate limiting for selenate uptake and assimilation.

Overexpression of ATP Sulfurylase in Indian Mustard Leads to Increased Selenate Uptake, Reduction, and Tolerance

Se can be accumulated by plants and volatilized to dimethylselenide, providing an attractive technology for Se phytoremediation. To determine the rate-limiting steps in Se volatilization from selenate and selenite, time- and concentration-dependent kinetics of Se accumulation and volatilization were studied in Indian mustard (Brassica juncea). Time-dependent kinetic studies showed that selenate was taken up 2-fold faster than selenite. Selenate was rapidly translocated to the shoot, away from the root, the site of volatilization, whereas only approximately 10% of the selenite was translocated. For both selenate- and selenite-supplied plants, Se accumulation and volatilization increased linearly with external Se concentration up to 20 mM; volatilization rates were also linearly correlated with root Se concentrations. Se-volatilization rates were 2- to 3-fold higher from plants supplied with selenite compared with selenate. Se speciation by x-ray absorption spectroscopy revealed that selenite-supplied plants accumulated organic Se, most likely selenomethionine, whereas selenate-supplied plants accumulated selenate. Our data suggest that Se volatilization from selenate is limited by the rate of selenate reduction, as well as by the availability of Se in roots, as influenced by uptake and translocation. Se volatilization from selenite may be limited by selenite uptake and by the conversion of selenomethionine to dimethylselenide.

Rate-Limiting Steps in Selenium Assimilation and Volatilization by Indian Mustard

Selenium is biologically important because (i) it is essential in animal and possibly plant metabolism, (ii) in many areas diets do not contain sufficient Se to meet, animals' needs, and (iii) in other areas it is toxic to animals when it occurrs in high concentrations in soil, water, plants, fly ash, or in aerosols. Animals require 0.05 to 0.1 mg Se/kg in their diets to prevent Se deficiency but suffer Se toxicosis when dietary levels exceed 5 to 15 mg Se/kg. The earth's crustal materials generally contain 50 mg Se/kg and more often contain <5 mg Se/kg. Soils and plants may discharge volatile forms of Se into the atmosphere. However, plants may also absorb measurable amounts of gaseous Se from the atmosphere. Anthropogenic activities impact the amount of Se entering our nation's lakes, rivers, and the atmosphere. Combustion of coal and incineration of municipal waste exhaust Se into the environment. In addition, crop-fallow and irrigation practices that allow leaching waters to pass through seleniferous strata prior to intersecting with surface flow, augment the Se levels encountered by plant and animal life.

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Selenium in Seleniferous Environments

An examination of the major anthropogenic and natural sources of atmospheric selenium indicates that approximately 13–19×109 g Se are cycled through the troposphere annually. Perhaps 60% of this flux is natural in origin, and both man-made and biological emissions appear to be predominantly gaseous in nature. Emissions from the marine biosphere and the combustion of coal account for 50–65% of the total atmospheric selenium emissions, and the natural marine flux appears to be roughly twice as large as that due to coal combustion.

A global atmospheric selenium budget

Samples collected from Kesterson Reservoir were screened for bacterial presence and selenate reduction capability. Selenate concentrations of 100 mg/liter were not toxic to indigenous bacteria. Of the 44 samples collected, 20 possessed microbial populations capable of reducing selenate. Reduction was observed in 4% of the water samples, 92% of the sediment samples, and 100% of the soil samples. Microbial reduction of 100 mg of selenate per liter was complete within 1 week of incubation. Up to 75 mg of selenate per liter was reduced beyond selenite to an insoluble red precipitate. Data collected indicate that indigenous bacteria have a significant role in the biogeochemical cycling of selenium.

R. Zieve

Papers

Volatilization of selenium from plants and soils