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28 MAYLAND ET AL. <br />Selenium-containing transfer ribonucleic acids (tRNA) have been found <br />in a number of aerobic and anaerobic bacteria, mammalian cells, and in Se- <br />accumulating and nonaccumulating plants (Wen et al., 1988). The Se- <br />containing tRNA appears to be ubiquitous in the plant kingdom. However, <br />the distribution of selenonucleoside(s) in tRNA species varies both qualita- <br />tively and quantitatively in different plants. This information provides evi- <br />dence for a possible biological role of Se in higher plants. Nevertheless, the <br />possibility that Se substituted for S in the nucleoside has not been evaluated. <br />The Se metabolites in plants are, generally, analogs of S compounds. <br />Nevertheless, Se metabolism in non-Se-accumulator plants cannot be identi- <br />fied from known mechanisms involving Se metabolism (Shrift, 1973). Our <br />understanding of the metabolic pathways for Se in plants remains very limited. <br />Many microorganisms can reduce selenite to elemental Se. Some bac- <br />teria and fungi have been found that reduce selenite or selenate to volatile, <br />organic Se compounds (Shrift, 1973). Sulfur-Se antagonism occurs in these <br />microorganisms. The reduction of selenite is enzymatically mediated, and <br />many isolated enzyme systems can utilize S and Se analogs interchangeably. <br />However, what was traditionally thought to be the mechanism of Se toxici- <br />ty, namely, a general interference with the enzymes involving S assimilation, <br />has proved to be more complex. <br />Selenium Toxicity in Plants <br />The toxicity of Se on the growth of grasses was noted as early as 1880 <br />(Brown & Shrift, 1982). However, the concern about selenosis in plants did <br />not really develop until the identification of selenosis in livestock and the <br />probable Se-associated reductions in wheat production in the seleniferous <br />areas (O.E. Olson, 1986 personal communication). <br />Selenium toxicity to nonaccumulator plants was described by Singh and <br />Singh (1978). They treated soil in the greenhouse with 0, 2.5, 5, or 10 mg <br />Se/kg as selenite and measured relative wheat-grain production of 100, 72, <br />17, and 11%, respectively. The grain contained 0.6, 10.2, 24.5, and 36.8 mg <br />Se/kg, respectively. The application of Se to the soil decreased the growth <br />of wheat and sunflower (Helianthus annuus L.); the absorption of Zn, Cu, <br />Fe, Mn, S, and N; and the synthesis of nucleic acids, chlorophyll, and pro- <br />tein, whereas additions of 50 or 100 mg P/kg as KzPO4 decreased the harm- <br />ful effect of Se and increased the values of each of the parameters tested. <br />Other plants, namely those classified by Rosenfeld and Beath (1964) as <br />primary or secondary indicator plants, tolerate and possibly require Se for <br />growth. These indicator plants differ from the nonaccumulator species in <br />several ways. Large quantities of Se-methylselenocysteine and selenocystathio- <br />nine, two nonprotein selenoamino acids rarely detected in nonaccumulators, <br />have been isolated from the tissues of Se accumulators (Brown & Shrift, 1982). <br />Also, Se is kept from entering proteins so that the Se level in proteins of <br />accumulator plants is significantly lower than the level in Se-sensitive plants. <br />Exclusion of Se from the proteins of accumulators is thought to be the basis <br />