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The characterization of ahyd gene cluster encoding the stable, bidirectional [NiFe]hydrogenase 1 enzyme inThiocapsa roseopersicina BBS, a purple sulfur photosynthetic bacterium belonging to the familyChromatiaceae , is presented. The heterodimeric hydrogenase 1 had been purified to homogeneity and thoroughly characterized (K. L. Kovacs et al., J. Biol. Chem. 266:947–951, 1991; C. Bagyinka et al., J. Am. Chem. Soc. 115:3567–3585, 1993). As an unusual feature, a 1,979-bp intergenic sequence (IS) separates the structural geneshydS andhydL , which encode the small and the large subunits, respectively. This IS harbors two sequential open reading frames (ORFs) which may code for electron transfer proteins ISP1 and ISP2. ISP1 and ISP2 are homologous to ORF5 and ORF6 in thehmc operon, coding for a transmembrane electron transfer complex inDesulfovibrio vulgaris . Other accessory proteins are not found immediately downstream or upstream ofhydSL . Ahup gene cluster coding for a typical hydrogen uptake [NiFe]hydrogenase inT. roseopersicina was reported earlier (A. Colbeau et al. Gene 140:25–31, 1994). The deduced amino acid sequences of the two small (hupS andhydS ) and large subunit (hupL andhydL ) sequences share 46 and 58% identity, respectively. Thehup andhyd genes differ in the arrangement of accessory genes, and the genes encoding the two enzymes are located at least 15 kb apart on the chromosome. Both hydrogenases are associated with the photosynthetic membrane. A stable and an unstable hydrogenase activity can be detected in cells grown under nitrogen-fixing conditions; the latter activity is missing in cells supplied with ammonia as the nitrogen source. The apparently constitutive and stable activity corresponds to hydrogenase 1, coded byhydSL , and the inducible and unstable second hydrogenase may be the product of thehup gene cluster.

Unusual Organization of the Genes Coding for HydSL, the Stable [NiFe]Hydrogenase in the Photosynthetic Bacterium Thiocapsa roseopersicina BBS