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The positions of the E. g. the Azotobacters). However, the E. coli Hyp proteins are consistently more deeply rooted and more closely match the relative positions of the hydrogenase 2 (hyb) genes rather than the hydrogenase 1 (hya) genes which group more closely with other ␥ group members. It is also interesting to note that the hyp genes in E. coli unlike those in other Proteobacteria cluster with the hyc (formate hydrogenase lyase) genes rather than the hya, hyb, or hyf genes. In E. coli, hypF is separate from the main hyp gene cluster (Maier et al.

The boxes show either the various groups of Eubacteria or genera of Archaea. No details are provided for the evolutionary relationships amongst the members of the Eukarya. Numbers at the ends of the branches represent the number of species of that Group or Genus in which hydrogenases or hydrogen metabolism has been studied or in which putative hydrogenase gene sequences have been discovered through genome sequence projects. 1. It should be noted that evolutionary distances are not to scale. g. Sulfolobus solfataricus, Pyrobaculum aerophilum and Aeropyrum pernix.

2C) with an apparently similar organisation of subunits and prosthetic groups to the Eubacterial examples of Group 5. Group 7. g. CO2) to CH4. Factor F420 (8-hydroxy 5-deazaflavin) is one of several unique cofactors required for methanogenesis (Fig. 2G). The two metalcontaining hydrogenases found in methanogens can be distinguished biochemically by their ability to reduce F420. The F420-non-reducing hydrogenases cannot reduce F420 but reduce the artificial electron acceptor MV (Deppenmeier et al.

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