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Asteroid and comet impacts are known to have caused profound disruption to multicellular life, yet their influence on habitats for microorganisms, which comprise the majority of Earth's biomass, is less well understood. Of particular interest are geological changes in the target lithology at and near the point of impact that can persist for billions of years. Deep subsurface and surface-dwelling microorganisms are shown to gain advantages from impact-induced fracturing of rocks. Deleterious changes are associated with impact-induced closure of pore spaces in rocks. Superimposed on these long-term geological changes are post-impact alterations such as changes in the hydrological system in and around a crater. The close coupling between geological changes and the conditions for microorganisms yields a synthesis of the fields of microbiology and impact cratering. We use these data to discuss how craters can be used in the search for life beyond Earth.

Unlabelled: The oxidation of molecular hydrogen (H2) is thought to be a major source of metabolic energy for life in the deep subsurface on Earth, and it could likewise support any extant biosphere on Mars, where stable habitable environments are probably limited to the subsurface. Faulting and fracturing may stimulate the supply of H2 from several sources. We report the H2 content of fluids present in terrestrial rocks formed by brittle fracturing on fault planes (pseudotachylites and cataclasites), along with protolith control samples. The fluids are dominated by water and include H2 at abundances sufficient to support hydrogenotrophic microorganisms, with strong H2 enrichments in the pseudotachylites compared to the controls. Weaker and less consistent H2 enrichments are observed in the cataclasites, which represent less intense seismic friction than the pseudotachylites. The enrichments agree quantitatively with previous experimental measurements of frictionally driven H2 formation during rock fracturing. We find that conservative estimates of current martian global seismicity predict episodic H2 generation by Marsquakes in quantities useful to hydrogenotrophs over a range of scales and recurrence times. On both Earth and Mars, secondary release of H2 may also accompany the breakdown of ancient fault rocks, which are particularly abundant in the pervasively fractured martian crust. This study strengthens the case for the astrobiological investigation of ancient martian fracture systems., Key Words: Deep biosphere-Faults-Fault rocks-Seismic activity-Hydrogen-Mars. Astrobiology 16, 690-702.

The interaction of microbes and metals is widely assumed to have occurred in surface or very shallow subsurface environments. However new evidence suggests that much microbial activity occurs in the deep subsurface. Fluvial, lacustrine and aeolian 'red beds' contain widespread centimetre-scale reduction spheroids in which a pale reduced spheroid in otherwise red rocks contains a metalliferous core. Most of the reduction of Fe (III) in sediments is caused by Fe (III) reducing bacteria. They have the potential to reduce a range of metals and metalloids, including V, Cu, Mo, U and Se, by substituting them for Fe (III) as electron acceptors, which are all elements common in reduction spheroids. The spheroidal morphology indicates that they were formed at depth, after compaction, which is consistent with a microbial formation. Given that the consequences of Fe (III) reduction have a visual expression, they are potential biosignatures during exploration of the terrestrial and extraterrestrial geological record. There is debate about the energy available from Fe (III) reduction on Mars, but the abundance of iron in Martian soils makes it one of the most valuable prospects for life there. Entrapment of the microbes themselves as fossils is possible, but a more realistic target during the exploration of Mars would be the colour contrasts reflecting selective reduction or oxidation. This can be achieved by analysing quartz grains across a reduction spheroid using Raman spectroscopy, which demonstrates its suitability for life detection in subsurface environments. Microbial action is the most suitable explanation for the formation of reduction spheroids and may act as metalliferous biosignatures for deep subsurface microbial activity.

There is abundant evidence for widespread microbial activity in deep continental fractures and aquifers, with important implications for biogeochemical cycling on Earth and the habitability of other planetary bodies. Whitman et al. (P Natl Acad Sci USA, 95, 1998, 6578) estimated a continental subsurface biomass on the order of 10(16) -10(17) g C. We reassess this value in the light of more recent data including over 100 microbial population density measurements from groundwater around the world. Making conservative assumptions about cell carbon content and the ratio of attached and free-living microorganisms, we find that the evidence continues to support a deep continental biomass estimate of 10(16) -10(17) g C, or 2-19% of Earth's total biomass., (© 2013 Federation of European Microbiological Societies. Published by John Wiley & Sons Ltd. All rights reserved.)

There is extensive evidence from drilling into continental margins for microbial colonization of a deep biosphere. However it is difficult to prove deep biosphere activity in the geological record, where evidence for life is dominated by the remains of organic matter buried after deposition at the surface. Nevertheless we propose that natural injections of sand into muddy strata at continental margins represent an excellent habitat opportunity for deep microbial activity down to several kilometres' present day depth. Sulphur isotope data for iron sulphides precipitated soon after injection indicate consistent microbial sulphate reduction through the geological record. The complexes are favourable sites for colonization, because high permeability and extensive sand/mud interface allow ready availability of electron donors and nutrients. The measured examples of iron sulphide in injected sands extend back to the Proterozoic, and show that injected sand complexes have been a long-term environment for deep subsurface microbial colonization.

Background: Horizontal gene transfer (HGT) plays a major role in speciation and evolution of bacteria and archaea by controlling gene distribution within an environment. However, information that links HGT to a natural community using relevant population-genetics parameters and spatial considerations is scarce. The Great Salt Lake (Utah, USA) provides an excellent model for studying HGT in the context of biogeography because it is a contiguous system with dispersal limitations due to a strong selective salinity gradient. We hypothesize that in spite of the barrier to phylogenetic dispersal, functional characteristics--in the form of HGT--expand beyond phylogenetic limitations due to selective pressure., Methodology and Results: To assay the functional genes and microorganisms throughout the GSL, we used a 16S rRNA oligonucleotide microarray (Phylochip) and a functional gene array (GeoChip) to measure biogeographic patterns of nine microbial communities. We found a significant difference in biogeography based on microarray analyses when comparing Sørensen similarity values for presence/absence of function and phylogeny (Student's t-test; p = 0.005)., Conclusion and Significance: Biogeographic patterns exhibit behavior associated with horizontal gene transfer in that informational genes (16S rRNA) have a lower similarity than functional genes, and functional similarity is positively correlated with lake-wide selective pressure. Specifically, high concentrations of chromium throughout GSL correspond to an average similarity of chromium resistance genes that is 22% higher than taxonomic similarity. This suggests active HGT may be measured at the population level in microbial communities and these biogeographic patterns may serve as a model to study bacteria adaptation and speciation.

The relationship between anthropogenic impact and the maintenance of biodiversity is a fundamental question in ecology. The emphasis on the organizational level of biodiversity responsible for ecosystem processes is shifting from a species-centred focus to include genotypic diversity. The relationship between biodiversity measures at these two scales remains largely unknown. By stratifying anthropogenic effects between scales of biodiversity of bacterial communities, we show a statistically significant difference in diversity based on taxonomic scale. Communities with intermediate species richness show high genotypic diversity while speciose and species-poor communities do not. We propose that in species-poor communities, generally comprising stable yet harsh conditions, physiological tolerance and competitive trade-offs limit both the number of species that occur and the loss of genotypes due to decreases in already constrained fitness. In species-rich communities, natural environmental conditions result in well-defined community structure and resource partitioning. Disturbance of these communities disrupts niche space, resulting in lower genotypic diversity despite the maintenance of species diversity. Our work provides a model to inform future research about relationships between species and genotypic biodiversity based on determining the biodiversity consequences of changing environmental context.