subsurface microbial community dominated by methanogens.
Nature, 415, 312–315.
Ciobanu, M.-C., Burgaude, G., Dufresne, A., Breuker, A., Redou,
V., Maamar, S. B., Gaboyer, F., Vandenabeele-Trambouze, O.,
Lipp, J., Schippers, A., Vandenkoornhuyse, P., Barbier, G.,
Jebbar, M., Godfroy, A., and Alaine, K., 2014. Microorganisms
persist at record-depths in the subseafloor of the Canterbury
basin. ISME Journal, 8, 1370–1380.
Coolen, M. J. L., Cypionka, H., Sass, A. M., Sass, H., and
Overmann, J., 2002. Ongoing modification of Mediterranean
Pleistocene sapropels mediated by prokaryotes. Science, 296,
2407–2410.
Coolen, M. J. L., Muyzer, G., Schouten, S., Volkman, J. K., and
Sinninghe Damsté, J. S., 2006. Sulfur and methane cycling during the Holocene in Ace Lake (Antarctica) revealed by lipid and
DNA stratigraphy. In Neretin, L. N. (ed.), Past and Present
Water Column Anoxia. Dordrecht: Springer. NATO Science
Series, IV. Earth and Environmental Sciences, Vol.
64, pp. 41–65.
Cowen, J. P., Giovannoni, S. J., Kenig, F., Johnson, H. P.,
Butterfield, D., Rappé, M. S., Hutnak, M., and Lam, P., 2003.
Fluids from aging ocean crust that support microbial life. Science, 299, 120–123.
Cragg, B. A., Parkes, R. J., Fry, J. C., Herbert, R. A., Wimpenny,
J. W. T., and Getliff, J. M., 1990. Bacterial biomass and activity
profiles within deep sediment layers. Proceedings of the Ocean
Drilling Program, Scientific Results, 112, 607–619.
Cragg, B. A., Parkes, R. J., Fry, J. C., Weightman, A. J., Rochelle,
P. A., and Maxwell, J. R., 1996. Bacterial populations and processes in sediments containing gas hydrates (ODP Leg 146:
Cascadia margin). Earth and Planetary Science Letters, 139,
497–507.
D’Hondt, S. L., Jørgensen, B. B., Miller, D. J., Batzke, A., Blake,
R., Cragg, B. A., Cypionka, H., Dickens, G. R., Ferdelman,
T. G., Hinrichs, K.-U., Holm, N. G., Mitterer, R., Spivack, A.,
Wang, G., Bekins, B., Engelen, B., Ford, K., Gettemy, G., Rutherford, S. D., Sass, H., Skilbeck, C. G., Aiello, I. W., Guèrin, G.,
House, C. H., Inagaki, F., Meister, P., Naehr, T., Niitsuma, S.,
Parkes, R. J., Schippers, A., Smith, D. C., Teske, A., Wiegel,
J., Padilla, C. N., and Solis Acosta, J. L., 2004. Distributions of
microbial activities in deep subseafloor sediments. Science,
306, 2216–2221.
D’Hondt, S., Spivack, A. J., Pockalny, R., Ferdelman, T. G.,
Fischer, J. P., Kallmeyer, J., Abrams, L. J., Smith, D. C., Graham,
D., Hasiuk, F., Schrum, H., and Stancin, A. M., 2009.
Subseafloor sedimentary life in the South Pacific gyre. Proceedings of the National Academy of Sciences of the United States of
America, 106, 11651–11656.
Durbin, A. M., and Teske, A., 2011. Microbial diversity and stratification of oligotrophic abyssal South Pacific sediments. Environmental Microbiology, 13, 3219–3234.
Durbin, A. M., and Teske, A., 2012. Archaea in organic-lean and
organic-rich marine subsurface sediments: an environmental
gradient reflected in distinct phylogenetic lineages. Frontiers in
Microbiology, 3, 168.
Edgcomb, V. P., Beaudoin, D., Gast, R., Biddle, J. F., and Teske, A.,
2011. Marine subsurface eukaryotes: the fungal majority. Environmental Microbiology, 13, 172–183.
Edwards, K. J., Bach, W., McCollom, T. M., and Rogers, D. R.,
2004. Neutrophilic iron-oxidizing bacteria in the ocean: their
habitats, diversity, and roles in mineral deposition, rock alteration, and biomass production in the deep-sea. Geomicrobiology
Journal, 21, 393–404.
Edwards, K. J., Bach, W., and McCollom, T. M., 2005.
Geomicrobiology in oceanography: mineral-microbe interactions at and below the seafloor. Trends in Microbiology, 13,
449–459.
Edwards, K. J., Becker, K., and Colwell, F., 2012. The deep, dark
energy biosphere: intraterrestrial life on Earth. Annual Review
of Earth and Planetary Sciences, 40, 551–568.
Engelen, B., Ziegelmüller, K., Wolf, L., Köpke, B., Gittel, A.,
Cypionka, H., Treude, T., Nakagawa, S., Inagaki, F., Lever,
M. A., and Steinsbu, B. O., 2008. Fluids from the oceanic crust
support microbial activities within the deep biosphere.
Geomicrobiology Journal, 25, 55–66.
Engelhardt, T., Sahlberg, M., Cypionka, H., and Engelen, B., 2013.
Biogeography of Rhizobium radiobacter and distribution of
associated temperate phages in deep subseafloor sediments.
ISME Journal, 7, 199–209.
Expedition 329 Scientists, 2011. South Pacific Gyre subseafloor
life. IODP Preliminary Report, 329, p. 108, doi:10.2204/iodp.
pr.329.2011.
Ferdelman, T. G., Fossing, H., Neumann, K., and Schulz, H. D.,
1999. Sulfate reduction in surface sediments of the southeast
Atlantic continental margin between 15
38
0 S and 27
57
0 S
(Angola and Namibia). Limnology and Oceanography, 44,
650–661.
Fisher, A. T., and Becker, K., 2000. Channelized fluid flow in oceanic crust reconciles heat-flow and permeability data. Nature,
403, 71–74.
Fisk, M. R., Giovannoni, S. J., and Furnes, H., 1998. Alteration of
oceanic volcanic glass textural evidence of microbial activity.
Science, 281, 978–980.
Fossing, H., Ferdelman, T. G., and Berg, P., 2000. Sulfate reduction
and methane oxidation in continental margin sediments
influenced by irrigation (South-East Atlantic off Namibia).
Geochimica et Cosmochimica Acta, 64, 897–910.
Fry, J. C., Parkes, R. J., Cragg, B. A., Weightman, A. J., and Webster, G., 2008. Prokaryotic diversity and activity in the deep
subseafloor biosphere. FEMS Microbiology Ecology, 66,
181–196.
Fry, J. C., Horsfield, B., Sykes, R., Cragg, B. A., Heywood, C.,
Kim, G. T., Mangelsdorf, K., Mildenhall, D. C., Rinna, J., Vieth,
A., Zink, K. G., Sass, H., Weightman, A. J., and Parkes, R. J.,
2009. Prokaryotic populations and activities in an interbedded
coal deposit, including a previously deeply buried section
(1.6–2.3 km) above $150 Ma basement rock. Geomicrobiology
Journal, 26, 163–178.
Gold, T., 1992. The deep, hot biosphere. Proceedings of the
National Academy of Sciences of the United States of America,
89, 6045–6049.
Gregg, W. W., Casey, N. W., and McClain, C. R., 2005. Recent
trends in global ocean chlorophyll. Geophysical Research Letters, doi:10.1029/2004gl021808.
Heberling, C., Lowell, R. P., Liu, L., and Fisk, M. R., 2010. Extent
of the microbial biosphere in the oceanic crust. Geochemistry,
Geophysics, Geosystems, 11(Q08003), 1–15.
Hedges, J. I., 1992. Global biogeochemical cycles: progress and
problems. Marine Chemistry, 39, 67–93.
Heid, C. A., Stevens, J., Livak, K. J., and Williams, P. M., 1996.
Real time quantitative PCR. Genome Research, 6, 986–994.
Heuer, V. B., Pohlman, J. W., Torres, M. E., Elvert, M., and
Hinrichs, K. U., 2009. The stable carbon isotope biogeochemistry of acetate and other dissolved carbon species in deep
subseafloor sediments at the northern Cascadia margin.
Geochimica et Cosmochimica Acta, 73, 3323–3336.
Hoehler, T. M., and Jørgensen, B. B., 2013. Microbial life under
extreme energy limitation. Nature Reviews Microbiology, 11,
83–94.
Hoehler, T. M., Alperin, M. J., Albert, D. B., and Martens, C. S.,
1994. Field and laboratory studies of methane oxidation in an
anoxic marine sediment: evidence for a methanogen-sulfate
reducer consortium. Global Biogeochemical Cycles, 8,
451–463.
152
DEEP BIOSPHERE
Nature, 415, 312–315.
Ciobanu, M.-C., Burgaude, G., Dufresne, A., Breuker, A., Redou,
V., Maamar, S. B., Gaboyer, F., Vandenabeele-Trambouze, O.,
Lipp, J., Schippers, A., Vandenkoornhuyse, P., Barbier, G.,
Jebbar, M., Godfroy, A., and Alaine, K., 2014. Microorganisms
persist at record-depths in the subseafloor of the Canterbury
basin. ISME Journal, 8, 1370–1380.
Coolen, M. J. L., Cypionka, H., Sass, A. M., Sass, H., and
Overmann, J., 2002. Ongoing modification of Mediterranean
Pleistocene sapropels mediated by prokaryotes. Science, 296,
2407–2410.
Coolen, M. J. L., Muyzer, G., Schouten, S., Volkman, J. K., and
Sinninghe Damsté, J. S., 2006. Sulfur and methane cycling during the Holocene in Ace Lake (Antarctica) revealed by lipid and
DNA stratigraphy. In Neretin, L. N. (ed.), Past and Present
Water Column Anoxia. Dordrecht: Springer. NATO Science
Series, IV. Earth and Environmental Sciences, Vol.
64, pp. 41–65.
Cowen, J. P., Giovannoni, S. J., Kenig, F., Johnson, H. P.,
Butterfield, D., Rappé, M. S., Hutnak, M., and Lam, P., 2003.
Fluids from aging ocean crust that support microbial life. Science, 299, 120–123.
Cragg, B. A., Parkes, R. J., Fry, J. C., Herbert, R. A., Wimpenny,
J. W. T., and Getliff, J. M., 1990. Bacterial biomass and activity
profiles within deep sediment layers. Proceedings of the Ocean
Drilling Program, Scientific Results, 112, 607–619.
Cragg, B. A., Parkes, R. J., Fry, J. C., Weightman, A. J., Rochelle,
P. A., and Maxwell, J. R., 1996. Bacterial populations and processes in sediments containing gas hydrates (ODP Leg 146:
Cascadia margin). Earth and Planetary Science Letters, 139,
497–507.
D’Hondt, S. L., Jørgensen, B. B., Miller, D. J., Batzke, A., Blake,
R., Cragg, B. A., Cypionka, H., Dickens, G. R., Ferdelman,
T. G., Hinrichs, K.-U., Holm, N. G., Mitterer, R., Spivack, A.,
Wang, G., Bekins, B., Engelen, B., Ford, K., Gettemy, G., Rutherford, S. D., Sass, H., Skilbeck, C. G., Aiello, I. W., Guèrin, G.,
House, C. H., Inagaki, F., Meister, P., Naehr, T., Niitsuma, S.,
Parkes, R. J., Schippers, A., Smith, D. C., Teske, A., Wiegel,
J., Padilla, C. N., and Solis Acosta, J. L., 2004. Distributions of
microbial activities in deep subseafloor sediments. Science,
306, 2216–2221.
D’Hondt, S., Spivack, A. J., Pockalny, R., Ferdelman, T. G.,
Fischer, J. P., Kallmeyer, J., Abrams, L. J., Smith, D. C., Graham,
D., Hasiuk, F., Schrum, H., and Stancin, A. M., 2009.
Subseafloor sedimentary life in the South Pacific gyre. Proceedings of the National Academy of Sciences of the United States of
America, 106, 11651–11656.
Durbin, A. M., and Teske, A., 2011. Microbial diversity and stratification of oligotrophic abyssal South Pacific sediments. Environmental Microbiology, 13, 3219–3234.
Durbin, A. M., and Teske, A., 2012. Archaea in organic-lean and
organic-rich marine subsurface sediments: an environmental
gradient reflected in distinct phylogenetic lineages. Frontiers in
Microbiology, 3, 168.
Edgcomb, V. P., Beaudoin, D., Gast, R., Biddle, J. F., and Teske, A.,
2011. Marine subsurface eukaryotes: the fungal majority. Environmental Microbiology, 13, 172–183.
Edwards, K. J., Bach, W., McCollom, T. M., and Rogers, D. R.,
2004. Neutrophilic iron-oxidizing bacteria in the ocean: their
habitats, diversity, and roles in mineral deposition, rock alteration, and biomass production in the deep-sea. Geomicrobiology
Journal, 21, 393–404.
Edwards, K. J., Bach, W., and McCollom, T. M., 2005.
Geomicrobiology in oceanography: mineral-microbe interactions at and below the seafloor. Trends in Microbiology, 13,
449–459.
Edwards, K. J., Becker, K., and Colwell, F., 2012. The deep, dark
energy biosphere: intraterrestrial life on Earth. Annual Review
of Earth and Planetary Sciences, 40, 551–568.
Engelen, B., Ziegelmüller, K., Wolf, L., Köpke, B., Gittel, A.,
Cypionka, H., Treude, T., Nakagawa, S., Inagaki, F., Lever,
M. A., and Steinsbu, B. O., 2008. Fluids from the oceanic crust
support microbial activities within the deep biosphere.
Geomicrobiology Journal, 25, 55–66.
Engelhardt, T., Sahlberg, M., Cypionka, H., and Engelen, B., 2013.
Biogeography of Rhizobium radiobacter and distribution of
associated temperate phages in deep subseafloor sediments.
ISME Journal, 7, 199–209.
Expedition 329 Scientists, 2011. South Pacific Gyre subseafloor
life. IODP Preliminary Report, 329, p. 108, doi:10.2204/iodp.
pr.329.2011.
Ferdelman, T. G., Fossing, H., Neumann, K., and Schulz, H. D.,
1999. Sulfate reduction in surface sediments of the southeast
Atlantic continental margin between 15
38
0 S and 27
57
0 S
(Angola and Namibia). Limnology and Oceanography, 44,
650–661.
Fisher, A. T., and Becker, K., 2000. Channelized fluid flow in oceanic crust reconciles heat-flow and permeability data. Nature,
403, 71–74.
Fisk, M. R., Giovannoni, S. J., and Furnes, H., 1998. Alteration of
oceanic volcanic glass textural evidence of microbial activity.
Science, 281, 978–980.
Fossing, H., Ferdelman, T. G., and Berg, P., 2000. Sulfate reduction
and methane oxidation in continental margin sediments
influenced by irrigation (South-East Atlantic off Namibia).
Geochimica et Cosmochimica Acta, 64, 897–910.
Fry, J. C., Parkes, R. J., Cragg, B. A., Weightman, A. J., and Webster, G., 2008. Prokaryotic diversity and activity in the deep
subseafloor biosphere. FEMS Microbiology Ecology, 66,
181–196.
Fry, J. C., Horsfield, B., Sykes, R., Cragg, B. A., Heywood, C.,
Kim, G. T., Mangelsdorf, K., Mildenhall, D. C., Rinna, J., Vieth,
A., Zink, K. G., Sass, H., Weightman, A. J., and Parkes, R. J.,
2009. Prokaryotic populations and activities in an interbedded
coal deposit, including a previously deeply buried section
(1.6–2.3 km) above $150 Ma basement rock. Geomicrobiology
Journal, 26, 163–178.
Gold, T., 1992. The deep, hot biosphere. Proceedings of the
National Academy of Sciences of the United States of America,
89, 6045–6049.
Gregg, W. W., Casey, N. W., and McClain, C. R., 2005. Recent
trends in global ocean chlorophyll. Geophysical Research Letters, doi:10.1029/2004gl021808.
Heberling, C., Lowell, R. P., Liu, L., and Fisk, M. R., 2010. Extent
of the microbial biosphere in the oceanic crust. Geochemistry,
Geophysics, Geosystems, 11(Q08003), 1–15.
Hedges, J. I., 1992. Global biogeochemical cycles: progress and
problems. Marine Chemistry, 39, 67–93.
Heid, C. A., Stevens, J., Livak, K. J., and Williams, P. M., 1996.
Real time quantitative PCR. Genome Research, 6, 986–994.
Heuer, V. B., Pohlman, J. W., Torres, M. E., Elvert, M., and
Hinrichs, K. U., 2009. The stable carbon isotope biogeochemistry of acetate and other dissolved carbon species in deep
subseafloor sediments at the northern Cascadia margin.
Geochimica et Cosmochimica Acta, 73, 3323–3336.
Hoehler, T. M., and Jørgensen, B. B., 2013. Microbial life under
extreme energy limitation. Nature Reviews Microbiology, 11,
83–94.
Hoehler, T. M., Alperin, M. J., Albert, D. B., and Martens, C. S.,
1994. Field and laboratory studies of methane oxidation in an
anoxic marine sediment: evidence for a methanogen-sulfate
reducer consortium. Global Biogeochemical Cycles, 8,
451–463.
152
DEEP BIOSPHERE