likely fed by hydrocarbons formed independently of photosynthesis in the ocean (Mason et al., 2010).
The size of the biomass in oceanic crust is unknown.
Ocean crust shall harbor the largest aquifer on Earth
(Fisher and Becker, 2000), which may transport molecular
oxygen and nitrate for basalt-weathering reactions
(Expedition 329 Scientists, 2011; Ziebis et al., 2012;
Orcutt et al., 2013b). This provides an enormous porous
space and chemical redox gradients as energy source for
a potentially huge deep biosphere. It has been suggested
based on model estimates that the size of the biomass of
the deep biosphere in oceanic crust is at least as big as that
in marine sediments (Heberling et al., 2010). Future
research will reveal this estimate.
Conclusions
The marine deep biosphere has been explored at various
ocean sites mainly within the framework of the Ocean
Drilling Program (ODP) and the Integrated Ocean Drilling
Program (IODP). The deep biosphere in the ocean crust
has been less intensively studied than that in subsurface
marine sediments. Unlike sediments in which organic carbon, derived from photosynthesis in the oceans, is the
major energy source for microbial life, inorganic processes in the ocean crusts likely provide the main energy
for the rock-hosted deep biosphere. The size of the biomass in oceanic crust is unknown. Kallmeyer et al. (2012)
estimated the values for the global biomass in
marine sediments being now 2.9 Â 10
29 cells,
corresponding to 4.1 petagram (Pg) C and ~0.6 % of
Earth’s total living biomass. The total microbial abundance in subseafloor sediments (2.9 Â 10
29 cells) is now
similar to the estimates for the total number of prokaryotes
in seawater (1.2 Â 10
29
) and in soil (2.6 Â 10
29
). The biomass of the deep subsurface comprises the three domains
of life Archaea, Bacteria, and Eukarya, as well as spores
and viruses. The analyses of 16S rRNA gene sequences
showed that microbial communities of deep marine sediments harbor members of distinct, uncultured bacterial
and archaeal phylogenetic lineages. Living cells, microbial activity, and the impact of the deep biosphere for biogeochemical processes have been demonstrated.
Bibliography
Amy, P. S., and Haldeman, D. L., 1997. The Microbiology of the
Terrestrial Deep Subsurface. New York: CRC.
Bach, W., and Edwards, K. J., 2003. Iron and sulfide oxidation
within the basaltic ocean crust: implications for chemolithoautotrophic microbial biomass production. Geochimica et
Cosmochimica Acta, 67, 3871–3887.
Bale, S. J., Goodman, K., Rochelle, P. A., Marchesi, J. R., Fry, J. C.,
Weightman, A. J., and Parkes, R. J., 1997. Desulfovibrio
profundus sp. nov., a novel barophilic sulfate-reducing bacteria
from deep sediments layers in the Japan Sea. International Journal of Systematic Bacteriology, 47, 515–521.
Barnes, S. P., Bradbrook, S. D., Cragg, B. A., Marchesi, J. R.,
Weightman, A. J., Fry, J. C., and Parkes, R. J., 1998. Isolation
of sulfate-reducing bacteria from deep sediment layers of the
Pacific Ocean. Geomicrobiology Journal, 15, 67–83.
Batzke, A., Engelen, B., Sass, H., and Cypionka, H., 2007. Phylogenetic and physiological diversity of cultured deep-biosphere bacteria from equatorial Pacific Ocean and Peru margin sediments.
Geomicrobiology Journal, 24, 261–273.
Biddle, J. F., House, C. H., and Brenchley, J. E., 2005.
Enrichment and cultivation of microorganisms from
sediment from the slope of the Peru Trench (ODP Site 1230).
In Jørgensen, B. B., D’Hondt, S. L., and Miller, D. J. (eds.), Proceedings of the ODP, Science Results, 201. Ocean Drilling Program, College Station, pp. 1–19. doi:10.2973/odp.proc.
sr.201.107.2005.
Biddle, J. F., Lipp, J. S., Lever, M., Lloyd, K. G., Sørensen, K. B.,
Anderson, R., Fredricks, H. F., Elvert, M., Kelly, T. J., Schrag,
D. P., Sogin, M. L., Brenchley, J. E., Teske, A., House, C. H.,
and Hinrichs, K.-U., 2006. Heterotrophic archaea dominate sedimentary subsurface ecosystems off Peru. Proceedings of the
National Academy of Sciences of the United States of America,
103, 3846–3851.
Biddle, J. F., Fitz-Gibbon, S., Schuster, S. C., Brenchley, J. E., and
House, C. H., 2008. Metagenomic signatures of the Peru margin
subseafloor biosphere show a genetically distinct environment.
Proceedings of the National Academy of Sciences of the United
States of America, 105, 10583–10588.
Biddle, J. F., White, J. R., Teske, A. P., and House, C. H., 2011.
Metagenomics of the subsurface Brazos-Trinity Basin (IODP
site 1320): comparison with other sediment and pyrosequenced
metagenomes. ISME Journal, 5, 1038–1047.
Blazejak, A., and Schippers, A., 2010. High abundance of JS-1- and
Chloroflexi-related Bacteria in deeply buried marine sediments
revealed by quantitative, real-time PCR. FEMS Microbiology
Ecology, 72, 198–207.
Blazejak, A., and Schippers, A., 2011. Real-time PCR quantification and diversity analysis of the functional genes aprA and dsrA
of sulfate-reducing prokaryotes in marine sediments of the Peru
continental margin and the Black Sea. Frontiers in Microbiology,
2, 253.
Boetius, A., Ravenschlag, K., Schubert, C. J., Rickert, D., Widdel,
F., Gieseke, A., Amann, R., Jorgensen, B. B., Witte, U., and
Pfannkuche, O., 2000. A marine microbial consortium apparently mediating anaerobic oxidation of methane. Nature, 407,
623–626.
Borgonie, G., García-Moyano, A., Litthauer, D., Bert, W., Bester,
A., van Heerden, E., Möller, C., Erasmus, M., and Onstott,
T. C., 2011. Nematoda from the terrestrial deep subsurface of
South Africa. Nature, 474, 79–82.
Borowski, W. S., Paull, C. K., and Ussler, W., III, 1997. Carbon
cycling with the upper methanogenic zone of continental rise
sediments: an example from the methane-rich sediments overlying the Blake Ridge gas hydrate deposits. Marine Geology, 157,
299–311.
Breuker, A., and Schippers, A., 2013. Data report: total cell counts
and qPCR abundance of Archaea and Bacteria in shallow subsurface marine sediments of North Pond: gravity cores collected
on site survey cruise prior to IODP Expedition 336. In Edwards,
K. J., Bach, W., Klaus, A., and Expedition 336 Scientists (eds.),
Proceedings of the IODP 336. Integrated Ocean Drilling Program Management International, Inc., Tokyo.
Breuker, A., Köweker, G., Blazejak, A., and Schippers, A., 2011.
The deep biosphere in terrestrial sediments of the Chesapeake
Bay impact structure, Virginia, USA. Frontiers in Microbiology,
2, 156.
Breuker, A., Stadler, S., and Schippers, A., 2013. Microbial community analysis of deeply buried marine sediments of the New
Jersey shallow shelf (IODP Expedition 313). FEMS Microbiology Ecology, 85, 578–592.
Chapelle, F. H., O’Neill, K., Bradley, P. M., Methé, B. A., Ciufo,
S. A., Knobel, L. L., and Lovley, D. R., 2002. A hydrogen-based
DEEP BIOSPHERE
151
The size of the biomass in oceanic crust is unknown.
Ocean crust shall harbor the largest aquifer on Earth
(Fisher and Becker, 2000), which may transport molecular
oxygen and nitrate for basalt-weathering reactions
(Expedition 329 Scientists, 2011; Ziebis et al., 2012;
Orcutt et al., 2013b). This provides an enormous porous
space and chemical redox gradients as energy source for
a potentially huge deep biosphere. It has been suggested
based on model estimates that the size of the biomass of
the deep biosphere in oceanic crust is at least as big as that
in marine sediments (Heberling et al., 2010). Future
research will reveal this estimate.
Conclusions
The marine deep biosphere has been explored at various
ocean sites mainly within the framework of the Ocean
Drilling Program (ODP) and the Integrated Ocean Drilling
Program (IODP). The deep biosphere in the ocean crust
has been less intensively studied than that in subsurface
marine sediments. Unlike sediments in which organic carbon, derived from photosynthesis in the oceans, is the
major energy source for microbial life, inorganic processes in the ocean crusts likely provide the main energy
for the rock-hosted deep biosphere. The size of the biomass in oceanic crust is unknown. Kallmeyer et al. (2012)
estimated the values for the global biomass in
marine sediments being now 2.9 Â 10
29 cells,
corresponding to 4.1 petagram (Pg) C and ~0.6 % of
Earth’s total living biomass. The total microbial abundance in subseafloor sediments (2.9 Â 10
29 cells) is now
similar to the estimates for the total number of prokaryotes
in seawater (1.2 Â 10
29
) and in soil (2.6 Â 10
29
). The biomass of the deep subsurface comprises the three domains
of life Archaea, Bacteria, and Eukarya, as well as spores
and viruses. The analyses of 16S rRNA gene sequences
showed that microbial communities of deep marine sediments harbor members of distinct, uncultured bacterial
and archaeal phylogenetic lineages. Living cells, microbial activity, and the impact of the deep biosphere for biogeochemical processes have been demonstrated.
Bibliography
Amy, P. S., and Haldeman, D. L., 1997. The Microbiology of the
Terrestrial Deep Subsurface. New York: CRC.
Bach, W., and Edwards, K. J., 2003. Iron and sulfide oxidation
within the basaltic ocean crust: implications for chemolithoautotrophic microbial biomass production. Geochimica et
Cosmochimica Acta, 67, 3871–3887.
Bale, S. J., Goodman, K., Rochelle, P. A., Marchesi, J. R., Fry, J. C.,
Weightman, A. J., and Parkes, R. J., 1997. Desulfovibrio
profundus sp. nov., a novel barophilic sulfate-reducing bacteria
from deep sediments layers in the Japan Sea. International Journal of Systematic Bacteriology, 47, 515–521.
Barnes, S. P., Bradbrook, S. D., Cragg, B. A., Marchesi, J. R.,
Weightman, A. J., Fry, J. C., and Parkes, R. J., 1998. Isolation
of sulfate-reducing bacteria from deep sediment layers of the
Pacific Ocean. Geomicrobiology Journal, 15, 67–83.
Batzke, A., Engelen, B., Sass, H., and Cypionka, H., 2007. Phylogenetic and physiological diversity of cultured deep-biosphere bacteria from equatorial Pacific Ocean and Peru margin sediments.
Geomicrobiology Journal, 24, 261–273.
Biddle, J. F., House, C. H., and Brenchley, J. E., 2005.
Enrichment and cultivation of microorganisms from
sediment from the slope of the Peru Trench (ODP Site 1230).
In Jørgensen, B. B., D’Hondt, S. L., and Miller, D. J. (eds.), Proceedings of the ODP, Science Results, 201. Ocean Drilling Program, College Station, pp. 1–19. doi:10.2973/odp.proc.
sr.201.107.2005.
Biddle, J. F., Lipp, J. S., Lever, M., Lloyd, K. G., Sørensen, K. B.,
Anderson, R., Fredricks, H. F., Elvert, M., Kelly, T. J., Schrag,
D. P., Sogin, M. L., Brenchley, J. E., Teske, A., House, C. H.,
and Hinrichs, K.-U., 2006. Heterotrophic archaea dominate sedimentary subsurface ecosystems off Peru. Proceedings of the
National Academy of Sciences of the United States of America,
103, 3846–3851.
Biddle, J. F., Fitz-Gibbon, S., Schuster, S. C., Brenchley, J. E., and
House, C. H., 2008. Metagenomic signatures of the Peru margin
subseafloor biosphere show a genetically distinct environment.
Proceedings of the National Academy of Sciences of the United
States of America, 105, 10583–10588.
Biddle, J. F., White, J. R., Teske, A. P., and House, C. H., 2011.
Metagenomics of the subsurface Brazos-Trinity Basin (IODP
site 1320): comparison with other sediment and pyrosequenced
metagenomes. ISME Journal, 5, 1038–1047.
Blazejak, A., and Schippers, A., 2010. High abundance of JS-1- and
Chloroflexi-related Bacteria in deeply buried marine sediments
revealed by quantitative, real-time PCR. FEMS Microbiology
Ecology, 72, 198–207.
Blazejak, A., and Schippers, A., 2011. Real-time PCR quantification and diversity analysis of the functional genes aprA and dsrA
of sulfate-reducing prokaryotes in marine sediments of the Peru
continental margin and the Black Sea. Frontiers in Microbiology,
2, 253.
Boetius, A., Ravenschlag, K., Schubert, C. J., Rickert, D., Widdel,
F., Gieseke, A., Amann, R., Jorgensen, B. B., Witte, U., and
Pfannkuche, O., 2000. A marine microbial consortium apparently mediating anaerobic oxidation of methane. Nature, 407,
623–626.
Borgonie, G., García-Moyano, A., Litthauer, D., Bert, W., Bester,
A., van Heerden, E., Möller, C., Erasmus, M., and Onstott,
T. C., 2011. Nematoda from the terrestrial deep subsurface of
South Africa. Nature, 474, 79–82.
Borowski, W. S., Paull, C. K., and Ussler, W., III, 1997. Carbon
cycling with the upper methanogenic zone of continental rise
sediments: an example from the methane-rich sediments overlying the Blake Ridge gas hydrate deposits. Marine Geology, 157,
299–311.
Breuker, A., and Schippers, A., 2013. Data report: total cell counts
and qPCR abundance of Archaea and Bacteria in shallow subsurface marine sediments of North Pond: gravity cores collected
on site survey cruise prior to IODP Expedition 336. In Edwards,
K. J., Bach, W., Klaus, A., and Expedition 336 Scientists (eds.),
Proceedings of the IODP 336. Integrated Ocean Drilling Program Management International, Inc., Tokyo.
Breuker, A., Köweker, G., Blazejak, A., and Schippers, A., 2011.
The deep biosphere in terrestrial sediments of the Chesapeake
Bay impact structure, Virginia, USA. Frontiers in Microbiology,
2, 156.
Breuker, A., Stadler, S., and Schippers, A., 2013. Microbial community analysis of deeply buried marine sediments of the New
Jersey shallow shelf (IODP Expedition 313). FEMS Microbiology Ecology, 85, 578–592.
Chapelle, F. H., O’Neill, K., Bradley, P. M., Methé, B. A., Ciufo,
S. A., Knobel, L. L., and Lovley, D. R., 2002. A hydrogen-based
DEEP BIOSPHERE
151