Three pathways for the specific incorporation of selenium into biological macromolecules. Selenophosphate is required for synthesis of selenocysteine (SeCys)-containing selenoproteins via cotranslational insertion of SeCys via selC tRNA that is first charged with serine. A specific elongation factor (SelB) interacts with the selenocysteine insertion sequence mRNA element located 3′ to the UGA codon (encoding the 21st amino acid, SeCys). This pathway was best defined in E. coli using the FDHH enzyme as the model (35, 42, 53). Selenophosphate is also required as a metabolic precursor for modification of uridine residues in several tRNAs (not SeCys tRNAs) to form 2-selenouridine. Both selenophosphate synthetase (SelD) and YbbB proteins have been shown to be required for this production of the selenouridine residue (63). In the third pathway, the incorporation of selenium into SDMHs in a labile form is not understood, and the dependence of this pathway on selenophosphate has not yet been established, although this labile form of selenium has been shown to occur in several molybdoenzymes isolated from clostridial species (14, 17, 18, 43-45, 49, 50). It has been proposed that this pathway exists in E. faecalis, based on two computational studies (20, 64), and that the first and second pathways are absent.