– Organization, heredity, growth, adaptation • But microbial life is distinctive: – Unicellular life allows niche exploitation of small space – Intimate association with environment – Well distributed through the habitable planet – Adapted to a range of temperatures, pressures, pH, salinities, and toxins – Numerous redox couples are exploited for energy – Reproduction by simple cell division – Long-term dormancy possible – Vast genetic diversity and exchange of information between individuals O’Dors Law of Biology: Physics and chemistry have laws, while biology only has lawyers looking for loopholes.
• Remarkable diversity of microbial life • Uncultured (i.e., unisolated) organisms are a major component • Large evolutionary divergence is present in the uncultured fraction • Huge physiological potential and activity apparently absent from biogeochemical models • Many of the uncultivated groups are in Earth environments
- Biomass decrease with depth - Numbers are lower than early estimates - Span 5 orders of magnitude - Strong correlation to mean sedimentation rate and distance from land (ocean productivity) Non-sedimentary crustal seafloor and continental settings progressing Fractures Priscu: “…vein volume in Antarctic continent is 144 km3” For seafloor sediments…
subsurface becomes too: - Hot (c.f., Roussel et al. 2008; highest temperature?) - Impermeable (Pedersen 2000; Colwell et al. 1997) - Dry (Colwell et al. 1992) - Compacted (Rebata-Landa and Santamarina, 2006) - Deprived of energy flux or supply of redox couples (Fredrickson et al. 1997; Røy et al. 2012) ….combined factors Endospores need to be considered Rebata-Landa and Santamarina, 2006
U G A C CC U A U A C G AC U G AG G C G G G A U C A A A A G U U U A U A A G U A A A G A A C C U G C U U A U G C G C U A U U G C C G A C C U G G U A A U G G C G A U G A C C C U A U A C G AC UG AG G C G G G A U C A A A A G U U U A U A A G U A A A G A A C C U G C U U A U G C G C U A U U G C C G A C C U G G U A A U Ribosomes in a microbial cell George Rice Ribosome A fragment of ribosomal RNA Central Dogma of Molecular Biology: “DNA makes RNA makes protein” F. Crick 1958 0.025 µm 1 x 5 µm 0.002 x 0.100 µm
2.8 km deep fracture Thermophilic Archaeal genes to fix N and C Uses H2 , CO >99.9% of the community Capable of independent existence Similar phylotypes detected in seafloor sediments
this study Rocks, sediments, waters Census of Deep Life Cratons, metamorphic rocks, basalts, sediments, near hydrothermal vents, karst caves, serpentinites, hydrates, springs, subglacial, sub ice shelf, BIFs, sapropels, tectonically active, sulfidic caves, mud volcanoes, seamounts
oxygen From Gaidos et al., 1999 Redox potential at 1 atm, pH 7, 25 C Reduced to oxidized half-reaction: energy rich (electrons to spare) to energy poor Oxidized to reduced half-reaction: energy poor (they crave electrons) to energy rich Physics and chemistry have laws, while biology only has lawyers looking for loopholes. Ron O’Dor Life needs a thermodynamic disequilibrium to survive
Onstott et al. 1999 • Mostly low activities in the subsurface; estimates by geochemical modeling • Subsurface rates range from 100 to 10-13 moles of CO2 /L/yr • Doubling times as low as 1013 sec; growth less typical than cell maintenance • Dependent upon balanced supply of electron donors and acceptors ….and many other factors
abundant, diverse, and metabolically tuned to long-term survival • Metabolic strategies for using redox couples are diverse and latent processes are likely more important than we have yet imagined • Understanding subsurface life as it is integrated with spatially and temporally expansive geological and chemical processes is an area of discovery