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microbial abundance and biomass in subseafloor sediment. Proc Natl Acad Sci U S A
109:16213–16216
Karnachuk OV, Frank YA, Lukina AP, Kadnikov VV, Beletsky AV, Mardanov AV, Ravin NV
(2019) Domestication of previously uncultivated Candidatus Desulforudis audaxviator from a
deep aquifer in Siberia sheds light on its physiology and evolution. ISME J 13(8):1947–1959.
https://doi.org/10.1038/s41396-019-0402-3
Kieft TL, McCuddy SM, Onstott TC, Davidson M, Lin L-H, Mislowac B, Pratt L, Boice E,
Sherwood Lollar B, Lippmann-Pipke J, Pfiffner SM, Phelps TJ, Gihring T, Moser D, van
Heerden A (2005) Geochemically generated, energy-rich substrates and indigenous microorganisms in deep, ancient groundwater. Geomicrobiol J 22:325–335
Kieft TL, Walters CC, Higgins MB, Mennito AS, Clewett CFM, Heuer V, Pullin MJ,
Hendrickson S, van Heerden E, Sherwood Lollar B, Lau MCY, Onstott TC (2018) Dissolved
organic matter compositions in 0.6–3.4 km deep fracture waters, Kaapvaal Craton, South Africa.
Org Geochem 118:116–131. https://doi.org/10.1016/j.orggeochem.2018.02.003
Lau MCY, Kieft TL, Kuloyo O, Linage-Alvarez B, van Heerden E, Lindsay MR, Magnabosco C,
Wang W, Wiggins JB, Guo L, Perlman DH, Kyin S, Shwe HH, Harris RL, Oh Y, Yi MJ,
Purtschert R, Slater GF, Ono S, Wei S, Li L, Lollar BS, Onstott TC (2016) Deep-subsurface
community dependent on syntrophy is dominated by sulfur-driven autotrophic denitrifiers. Proc
Natl Acad Sci U S A 113:E7927–E7936. https://doi.org/10.1073/pnas.1612244113
Lin LH, Wang P-L, Rumble D, Lippmann-Pipke J, Boice E, Pratt LM, Sherwood Lollar B,
Brodie E, Hazen T, Andersen G, DeSantis T, Moser DP, Kershaw D, Onstott TC (2006)
Long term biosustainability in a high energy, low diversity crustal biome. Science 314:479–482
Lippmann-Pipke J, Erzinger J, Zimmer M, Kujawa C, Boettcher M, van Heerde E, Bester A,
Moller H, Stroncik NA, Reches Z (2011) Geogas transport in fractured hard rock—correlations
with mining seismicity at 3.54 km depth, TauTona gold mine South Africa. Appl Geochem
26:2134–2146. https://doi.org/10.1016/j.apgeochem.2011.07.011
Locey KJ, Lennon JT (2016) Scaling laws predict global microbial diversity. Proc Natl Acad Sci U
S A 113(21):5970–5975. https://doi.org/10.1073/pnas.1521291113
Magnabosco C, Lin L-H, Dong H, Bomberg M, Ghiorse W, Stan-Lotter H, Pedersen K, Kieft TL,
vanHeerden E, Onstott TC (2018) The biomass and biodiversity of the continental subsurface.
Nat Geosci 11:707–717. https://doi.org/10.1038/s41561-018-0221-6
Nealson KH, Inagaki F, Takai K (2005) Hydrogen-driven subsurface lithoautotrophic microbial
ecosystems (SLiMEs): do they exist and why should we care? Trends in Microbiol 13:405–410.
https://doi.org/10.1016/j.tim.2005.07.010
Onstott TC, Ehlmann BL, Sapers H, Coleman M, Ivarsson M, Marlow JJ, Neubeck A, Niles P
(2018) Paleo-rock-hosted life on earth and the search on Mars: a review and strategy for
exploration. Astrobiology 19(10):1230–1261. https://doi.org/10.1089/ast.2018.1960
Onstott TC, Magnabosco C, Aubrey AD, Burton AS, Dworkin JP, Elsila JE, Grunsfeld S, Cao BH,
Hein JE, Glavin DP, Kieft TL, Silver BJ, Phelps TJ, van Heerden E, Opperman DJ, Bada JL
(2014) Does aspartic acid racemization constrain the depth limit of the subsurface biosphere?
Geobiology 12:1–19
Pedersen K (2000) Exploration of deep intraterrestrial microbial life: current perspectives. FEMS
Microbiol Lett 185:9–16
Schrenk MO, Brazelton WJ, Lang SQ (2013) Serpentinization, carbon and deep life. Rev Mineral
Geochem 75:575–606
Sherwood Lollar B, Onstott TC, Lacrampe-Couloume G, Ballantine CJ (2014) The contribution of
the Precambrian continental lithosphere to global H 2 production. Nature 516:379–382
Stevens TO, McKinley JP (1995) Lithoautotrophic microbial ecosystems in deep basaltic aquifers.
Science 270(5235):450–454. https://doi.org/10.1126/science.270.5235.450
Stevens TO, McKinley JP (2000) Abiotic controls on H2 production from basalt-water reactions
and implications for aquifer biogeochemistry. Environ Sci Technol 34:826–831
138
T. L. Kieft
microbial abundance and biomass in subseafloor sediment. Proc Natl Acad Sci U S A
109:16213–16216
Karnachuk OV, Frank YA, Lukina AP, Kadnikov VV, Beletsky AV, Mardanov AV, Ravin NV
(2019) Domestication of previously uncultivated Candidatus Desulforudis audaxviator from a
deep aquifer in Siberia sheds light on its physiology and evolution. ISME J 13(8):1947–1959.
https://doi.org/10.1038/s41396-019-0402-3
Kieft TL, McCuddy SM, Onstott TC, Davidson M, Lin L-H, Mislowac B, Pratt L, Boice E,
Sherwood Lollar B, Lippmann-Pipke J, Pfiffner SM, Phelps TJ, Gihring T, Moser D, van
Heerden A (2005) Geochemically generated, energy-rich substrates and indigenous microorganisms in deep, ancient groundwater. Geomicrobiol J 22:325–335
Kieft TL, Walters CC, Higgins MB, Mennito AS, Clewett CFM, Heuer V, Pullin MJ,
Hendrickson S, van Heerden E, Sherwood Lollar B, Lau MCY, Onstott TC (2018) Dissolved
organic matter compositions in 0.6–3.4 km deep fracture waters, Kaapvaal Craton, South Africa.
Org Geochem 118:116–131. https://doi.org/10.1016/j.orggeochem.2018.02.003
Lau MCY, Kieft TL, Kuloyo O, Linage-Alvarez B, van Heerden E, Lindsay MR, Magnabosco C,
Wang W, Wiggins JB, Guo L, Perlman DH, Kyin S, Shwe HH, Harris RL, Oh Y, Yi MJ,
Purtschert R, Slater GF, Ono S, Wei S, Li L, Lollar BS, Onstott TC (2016) Deep-subsurface
community dependent on syntrophy is dominated by sulfur-driven autotrophic denitrifiers. Proc
Natl Acad Sci U S A 113:E7927–E7936. https://doi.org/10.1073/pnas.1612244113
Lin LH, Wang P-L, Rumble D, Lippmann-Pipke J, Boice E, Pratt LM, Sherwood Lollar B,
Brodie E, Hazen T, Andersen G, DeSantis T, Moser DP, Kershaw D, Onstott TC (2006)
Long term biosustainability in a high energy, low diversity crustal biome. Science 314:479–482
Lippmann-Pipke J, Erzinger J, Zimmer M, Kujawa C, Boettcher M, van Heerde E, Bester A,
Moller H, Stroncik NA, Reches Z (2011) Geogas transport in fractured hard rock—correlations
with mining seismicity at 3.54 km depth, TauTona gold mine South Africa. Appl Geochem
26:2134–2146. https://doi.org/10.1016/j.apgeochem.2011.07.011
Locey KJ, Lennon JT (2016) Scaling laws predict global microbial diversity. Proc Natl Acad Sci U
S A 113(21):5970–5975. https://doi.org/10.1073/pnas.1521291113
Magnabosco C, Lin L-H, Dong H, Bomberg M, Ghiorse W, Stan-Lotter H, Pedersen K, Kieft TL,
vanHeerden E, Onstott TC (2018) The biomass and biodiversity of the continental subsurface.
Nat Geosci 11:707–717. https://doi.org/10.1038/s41561-018-0221-6
Nealson KH, Inagaki F, Takai K (2005) Hydrogen-driven subsurface lithoautotrophic microbial
ecosystems (SLiMEs): do they exist and why should we care? Trends in Microbiol 13:405–410.
https://doi.org/10.1016/j.tim.2005.07.010
Onstott TC, Ehlmann BL, Sapers H, Coleman M, Ivarsson M, Marlow JJ, Neubeck A, Niles P
(2018) Paleo-rock-hosted life on earth and the search on Mars: a review and strategy for
exploration. Astrobiology 19(10):1230–1261. https://doi.org/10.1089/ast.2018.1960
Onstott TC, Magnabosco C, Aubrey AD, Burton AS, Dworkin JP, Elsila JE, Grunsfeld S, Cao BH,
Hein JE, Glavin DP, Kieft TL, Silver BJ, Phelps TJ, van Heerden E, Opperman DJ, Bada JL
(2014) Does aspartic acid racemization constrain the depth limit of the subsurface biosphere?
Geobiology 12:1–19
Pedersen K (2000) Exploration of deep intraterrestrial microbial life: current perspectives. FEMS
Microbiol Lett 185:9–16
Schrenk MO, Brazelton WJ, Lang SQ (2013) Serpentinization, carbon and deep life. Rev Mineral
Geochem 75:575–606
Sherwood Lollar B, Onstott TC, Lacrampe-Couloume G, Ballantine CJ (2014) The contribution of
the Precambrian continental lithosphere to global H 2 production. Nature 516:379–382
Stevens TO, McKinley JP (1995) Lithoautotrophic microbial ecosystems in deep basaltic aquifers.
Science 270(5235):450–454. https://doi.org/10.1126/science.270.5235.450
Stevens TO, McKinley JP (2000) Abiotic controls on H2 production from basalt-water reactions
and implications for aquifer biogeochemistry. Environ Sci Technol 34:826–831
138
T. L. Kieft