Various classes of microbial and biomolecular evidence from global studies in marine and continental settings are used to identify a set of reactions that appear to support microbial metabolism during serpentinization of ultramafic rocks. Geochemical data from serpentinizing ecosystems in the Samail ophiolite of Oman are used to evaluate the extent of disequilibria that can support this set of microbial metabolisms and to provide a ranking of potential metabolic energy sources in hyperalkaline fluids that are direct products of serpentinization. Results are used to construct hypotheses for how microbial metabolism may be supported in the subsurface for two cases: ecosystems hosted in rocks that have already undergone significant serpentinization and those hosted by deeper, active serpentinization processes.

Calculations are presented of the extent to which chemical disequilibria are generated when submarine hydrothermal fluids mix with sea water. These calculations involve quantifying the chemical affinity for individual reactions by comparing equilibrium compositions with the compositions of mixtures in which oxidation-reduction reactions are inhibited. The oxidation-reduction reactions that depart from equilibrium in these systems provide energy for chemotrophic microbial metabolism. Methanogenesis is an example of this phenomenon, in which the combination of carbon dioxide, hydrogen and methane induced by fluid mixing is far from equilibrium, which can be approached if more methane is generated. Similarly, the production of other organic compounds is also favorable under the same conditions that permit methanogenesis. Alkanes, alkenes, alcohols, aldehydes, carboxylic acids and amino acids are among the compounds that, if formed, would lower the energetic state of the chemical composition generated in mixed fluids. The resulting positive values of chemical affinity correspond to the thermodynamic drive required for abiotic organic synthesis. It is also possible that energy release accompanies biosynthesis by chemotrophic organisms. In this way, hydrothermal ecosystems differ radically from familiar ecosystems at Earth's surface. If captured, the energy released may be sufficient to drive biosynthesis of carbohydrates, purines, pyrimidines and other compounds which require energy inputs.

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Serpentinizing systems, established when aqueous planetary fluids react with ultramafic (UM) rocks, produce significant quantities of dihydrogen (H 2 ) across a large range of geochemical conditions. This H 2 actively supports chemosynthetic communities on Earth (

Thermodynamic characterization of the relative stabilities of chemical compounds is a pillar of conceptual models in various fields of geosciences. Analogous models applied to genomes can yield new information about the relationship between genomes and their geochemical environments. In this perspective article, we present a chemical and thermodynamic analysis of prokaryotic lineages that have been the target of previous phylogenomic studies of evolutionary adaptation to varying redox conditions. The thermodynamic model development begins by quantifying the effects of hydrogen activity (aH2) and temperature on the relative stabilities of organic compounds with different carbon oxidation state. When applied to proteins instead of metabolites, the same techniques can be used to identify combinations of aH2 and temperature at which reference proteomes for Class I or Class II methanogens are relatively stable. The calculated aH2 values are compatible with reported measurements for habitats of methanogens ranging from highly reducing submarine hydrothermal systems to less reducing environments including methanogenic sediments. In contrast to the transition between the two classes of methanogenic archaea, that between basal and terrestrial groups of Thaumarchaeota (denoting the origin of ammonia‐oxidizing archaea) occurs at a less‐reducing redox boundary. These examples reveal the consequences of energy minimization driving evolution and show how geochemical calculations involving biomolecules can be used to quantify and better understand the coevolution of the geosphere and biosphere.

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