coli enzyme [90]), which are missing in G. metallireducens. Thus, G. sulfurreducens is capable of achieving osmotolerance Talazoparib without consuming carbohydrate storage polymers, but G. metallireducens is not. Biogenesis of c-type cytochromes and pili The genome of G. metallireducens encodes 91 putative c-type cytochromes, of which 65 have homologs among the 103 c-type cytochromes of G. sulfurreducens. Of the c-type cytochrome genes implicated in Fe(III) and U(VI) reduction in G. sulfurreducens, those conserved in G. metallireducens are macA (Gmet_3091 = GSU0466) [91–93] and ppcA (Gmet_2902 = GSU0612) [37], whereas different c-type cytochrome sequences are found in syntenous locations
where one would expect omcB and omcC (Gmet_0910 ≠ GSU2737; Gmet_0913 ≠ GSU2731) [94], and omcE (Gmet_2896 ≠ GSU0618) [95]. The G. metallireducens genome contains no genes homologous to omcS (GSU2504) and omcT (GSU2503) [95], and only a paralog (Gmet_0155 = GSU2743) of omcF (GSU2432) [96]. This lack of conservation is being investigated GDC-0449 research buy further (J. Butler, personal communication). Notable differences between G. metallireducens and G. sulfurreducens are apparent in the biogenesis of c-type cytochromes, in biosynthesis of the heme group, and in reduction of disulfide bonds to allow covalent linkage to heme. In addition
to the membrane-peripheral protoporphyrinogen IX oxidase of G. sulfurreducens and other Geobacteraceae, encoded by the hemY gene (Gmet_3551 = GSU0012, 38% identical to the Myxococcus xanthus enzyme [97]), G. metallireducens
has a membrane-integral isoenzyme encoded by hemG (Gmet_2953, 43% identical to the E. coli enzyme [98]), with a homolog in Geobacter FRC-32. These two species also possess a putative disulfide bond reduction system not found in G. sulfurreducens and other Geobacteraceae, comprised of DsbA, DsbB, DsbE and DsbD homologs (Gmet_1380, Gmet_1381, Gmet_1383, Gmet_1384), encoded in a cluster alongside a two-component signalling system (Gmet_1378-Gmet_1379), an arylsulfotransferase Y-27632 2HCl (Gmet_1382), and a conserved protein of unknown function (Gmet_1385). Transcription of dsbA and dsbB is diminished during growth on benzoate [21], and phylogenetic analysis indicates that these DsbA and DsbB proteins belong to subfamilies distinct from those that have been characterized (R. Dutton, personal communication). Located apart from this cluster, DsbC/DsbG (Gmet_2250) of G. metallireducens has homologs in several Geobacteraceae, but not in G. sulfurreducens. However, CcdA/DsbD (Gmet_2451 = GSU1322) is present in both. Thus, the pathways of c-type cytochrome biogenesis may be significantly different in the two species and somehow linked to the degradation of aromatic compounds by G. metallireducens. In both G. sulfurreducens and G. metallireducens, there are four c-type cytochrome biogenesis genes related to ResB of B.