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Submarine hydrothermal vents are model systems for the Archaean Earth environment, and some sites maintain conditions that may have favored the formation and evolution of cellular life. Vents are typified by rapid fluctuations in temperature and redox potential that impose a strong selective pressure on resident microbial communities.  Nautilia profundicola  strain Am-H is a moderately thermophilic, deeply-branching  Epsilonproteobacterium  found free-living at hydrothermal vents and is a member of the microbial mass on the dorsal surface of vent polychaete,  Alvinella pompejana . Analysis of the 1.7-Mbp genome of  N. profundicola  uncovered adaptations to the vent environment—some unique and some shared with other  Epsilonproteobacterial  genomes. The major findings included: (1) a diverse suite of hydrogenases coupled to a relatively simple electron transport chain, (2) numerous stress response systems, (3) a novel predicted nitrate assimilation pathway with hydroxylamine as a key intermediate, and (4) a gene ( rgy ) encoding the hallmark protein for hyperthermophilic growth, reverse gyrase. Additional experiments indicated that expression of  rgy  in strain Am-H was induced over 100-fold with a 20°C increase above the optimal growth temperature of this bacterium and that closely related  rgy  genes are present and expressed in bacterial communities residing in geographically distinct thermophilic environments.  N. profundicola , therefore, is a model  Epsilonproteobacterium  that contains all the genes necessary for life in the extreme conditions widely believed to reflect those in the Archaean biosphere—anaerobic, sulfur, H 2 - and CO 2 -rich, with fluctuating redox potentials and temperatures. In addition, reverse gyrase appears to be an important and common adaptation for mesophiles and moderate thermophiles that inhabit ecological niches characterized by rapid and frequent temperature fluctuations and, as such, can no longer be considered a unique feature of hyperthermophiles.

Adaptations to Submarine Hydrothermal Environments Exemplified by the Genome of Nautilia profundicola