All of the sequences were retrieved from SILVA [60] when available or GenBank (http://www.ncbi.nlm.nih.gov/). SOR are represented with a single green arrow, Dx-SOR with a double khaki arrow, Fe-Mn SOD by YH25448 in vivo a light blue dot
and Cu-Zn SOD by a dark blue dot. SOD-type genes were determined using OxyGene [36]. Scale bar: 3% difference. Crenarchaeota (in red) are developed in Figure 4. Nanoarchaeota [61] and Korarchaeota [62] are obligately anaerobic sulphur-dependent organisms placed close to the root of the archaeal SSU rRNA tree. Nanoarchaeota is currently known from a single organism Candidatus Nanoarchaeum equitans, a hyperthermophilic symbiont that grows on the surface of Ignicoccus hospitalis [62, 63]. There are currently no representatives of Korarchaeota in pure culture but the genome of K. cryptophilum, a very thin filamentous thermophilic heterotroph, has been determined from a sample of Yellowstone National Park TEW-7197 chemical structure Obsidian Pool. Both C. N. equitans and K. cryptophilum are found together in the 16S tree, in the vicinity of the Crenarchaeota group, and contain genes encoding superoxide reductase
AZD6094 in vivo with a SOR (centre II) functional domain and do not encode superoxide dismutase genes. According to 16S rRNA gene sequences, the Crenarchaeota group can be subdivided into three orders, the Thermoproteales, the Sulfolobales and the Desulfurococcales [64]. All Sulfolobales and Thermoproteoles genomes studied encode a single SOD, with the single exception of the unique member of the Thermofilaceae familly, Thermofilum pendens, an anaerobic commensal that encodes a SOR. By contrast, all Desulfurococcales genomes available encode a SOR but not a SOD, except Aeropyrum pernix that has the particularity to be strictly aerobic [65] and that encodes an extremely thermostable Mn/Fe superoxide dismutase [66] and Ignisphaera aggregans, a novel deep-branching member of the Desulfurococcaceae lineage of strict anaerobes (as even trace quantities of oxygen inhibited its
growth, [67] ) the genome Suplatast tosilate of which carries neither SOR or SOD genes. Other Desulfurococcales studied (Figure 4) have all a gene encoding a centre II mono-domain SOR-type enzyme. Interestingly, two recent genomes have been made available since the last update of SORGOdb (May 2010) and both contain annotation for SOR-like genes: Tagg_0590, described as a Desulfoferrodoxin ferrous iron-binding protein of Thermosphaera aggregans DSM 11486 and Shell_0770 for Staphylothermus hellenicus DSM 12710, annotated as a twin-arginine secreted superoxide reductase, by homology with Geobacter metallireducens GS-15 Gmet_2613 SOR. Using the SORGOdb “”search by BlastP”", we could confirm that both ORFs are true SOR (ten best e-value from e-59 to e-34) and belong to the SOR-type class.