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Prokaryotes: The unseen majority
Open accessAbstract
The number of prokaryotes and the total amount of their cellular carbon on earth are estimated to be 4–6 × 10 30 cells and 350–550 Pg of C (1 Pg = 10 15 g), respectively. Thus, the total amount of prokaryotic carbon is 60–100% of the estimated total carbon in plants, and inclusion of prokaryotic carbon in global models will almost double estimates of the amount of carbon stored in living organisms. In addition, the earth’s prokaryotes contain 85–130 Pg of N and 9–14 Pg of P, or about 10-fold more of these nutrients than do plants, and represent the largest pool of these nutrients in living organisms. Most of the earth’s prokaryotes occur in the open ocean, in soil, and in oceanic and terrestrial subsurfaces, where the numbers of cells are 1.2 × 10 29 , 2.6 × 10 29 , 3.5 × 10 30 , and 0.25–2.5 × 10 30 , respectively. The numbers of heterotrophic prokaryotes in the upper 200 m of the open ocean, the ocean below 200 m, and soil are consistent with average turnover times of 6–25 days, 0.8 yr, and 2.5 yr, respectively. Although subject to a great deal of uncertainty, the estimate for the average turnover time of prokaryotes in the subsurface is on the order of 1–2 × 10 3 yr. The cellular production rate for all prokaryotes on earth is estimated at 1.7 × 10 30 cells/yr and is highest in the open ocean. The large population size and rapid growth of prokaryotes provides an enormous capacity for genetic diversity.
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“However, the majority of astrobiological research is focused on the possible existence and detection of microbial life, which is the simplest, most robust, and most abundant form of life on our own planet (Whitman et al., 1998) and therefore is considered to be the most likely form of life to exist on other planets and moons.”
From The Astrobiology Primer v2.0 (2016) · cites this paper as Whitman et al., 1998
In the citing paper
“Given that there are estimated to be greater than 1030 prokaryotes on earth (Whitman et al., 1998), the vast majority of which are yet unstudied, there are likely to be more PFAS‐metabolizing bacteria uncovered.”
From Nothing lasts forever: understanding microbial biodegradation of polyfluorinated compounds and perfluorinated alkyl substances (2022) · cites this paper as Whitman et al., 1998
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Microbial view of central nervous system autoimmunity
Berer, K., & Krishnamoorthy, G. (2014) · FEBS Letters · doi:10.1016/j.febslet.2014.04.007 · Cited by 153
Fredriksson, N. J., Hermansson, M., & Wilén, B. (2013) · PLoS ONE · doi:10.1371/journal.pone.0076431 · Cited by 156
“For example, estimates for bacterial communities in soil are in the range of 107 -1010 bacterial cells [1,2] of 103 -105 different taxa [2,3].”
BackgroundThe Physiology of Phagocytosis in the Context of Mitochondrial Origin
Martin, W. F., Tielens, A. G. M., Mentel, M., Garg, S. G., & Gould, S. B. (2017) · Microbiology and Molecular Biology Reviews · doi:10.1128/mmbr.00008-17 · Cited by 157
“(319) estimated that roughly 1030 prokaryotic cells exist on Earth today.”
BackgroundMacroecology to Unite All Life, Large and Small
Shade, A. L., Dunn, R. R., Blowes, S. A., Keil, P., Bohannan, B. J. M., Herrmann, M., Küsel, K., Lennon, J. T., Sanders, N. J., Štorch, D., et al. (2018) · Trends in Ecology & Evolution · doi:10.1016/j.tree.2018.08.005 · Cited by 156
Suttle, C. A. (2000) · Elsevier eBooks · doi:10.1016/b978-012362675-2/50007-0 · Cited by 150
Priester, J. H., Stoimenov, P. K., Mielke, R. E., Webb, S. M., Ehrhardt, C. J., Zhang, J. P., Stucky, G. D., & Holden, P. A. (2009) · Environmental Science & Technology · doi:10.1021/es802806n · Cited by 151
Soil bacterial diversity mediated by microscale aqueous-phase processes across biomes
Bickel, S., & Or, D. (2020) · Nature Communications · doi:10.1038/s41467-019-13966-w · Cited by 150
Multicellular microorganisms: laboratory versus nature
Palková, Z. (2004) · EMBO Reports · doi:10.1038/sj.embor.7400145 · Cited by 151
“…density and variability of microorganisms that exist in different environments indicate that they represent one-half of all cellular carbon, but that 99% of them are unknown and cannot be grown under standard laboratory conditions (Whitman et al, 1998); in other words, only a small proportion (1”
BackgroundDell’Anno, A., & Corinaldesi, C. (2004) · Applied and Environmental Microbiology · doi:10.1128/aem.70.7.4384-4386.2004 · Cited by 150
“In order to explore the potential ecological implication of extracellular DNA in benthic biogeochemical processes, we estimated the daily bacterial P requirement on the basis of bacterial C production (assuming a bacterial C/N/P ratio of 40:10:1 [18]) and the daily P supply derived from DNA degradation rates (assuming…”
BackgroundProkaryote diversity and taxonomy: current status and future challenges
Oren, A. (2004) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2003.1458 · Cited by 151
“With an estimated total biomass of 350–550 × 10(15) g of carbon, prokaryotic cells contain as much as 60–100% of the estimated total carbon in plants (including extracellular material such as cell walls and structural polymers; Whitman et al. 1998).”
BackgroundInfluentialBarton, H. A., Taylor, N. M., Lübbers, B., & Pemberton, A. C. (2005) · Journal of Microbiological Methods · doi:10.1016/j.mimet.2005.10.005 · Cited by 159
Desulfotomaculum and Methanobacterium spp. Dominate a 4- to 5-Kilometer-Deep Fault
Moser, D. P., Gihring, T. M., Brockman, F. J., Fredrickson, J. K., Balkwill, D. L., Dollhopf, M. E., Lollar, B. S., Pratt, L. M., Boice, E., Southam, G., et al. (2005) · Applied and Environmental Microbiology · doi:10.1128/aem.71.12.8773-8783.2005 · Cited by 177
“, liquid water, habitable space, and permissive temperatures), it follows that a significant proportion of the biosphere may be microbial and associated with deep terrestrial hydrologic systems (76).”
BackgroundModelling the emergent dynamics and major metabolites of the human colonic microbiota
Kettle, H., Louis, P., Holtrop, G., Duncan, S. H., & Flint, H. J. (2014) · Environmental Microbiology · doi:10.1111/1462-2920.12599 · Cited by 149
“The human colon is one of the most densely populated microbial ecosystems on Earth (Whitman et al., 1998) with several hundred different bacterial strains coexisting at any one time (Eckburg et al.”
BackgroundThe microbial contribution to macroecology
Barberán, A. (2014) · Frontiers in Microbiology · doi:10.3389/fmicb.2014.00203 · Cited by 149
“First, microbial communities could expand the number of species and individuals included in macroecological datasets of plant and animal communities by several orders of magnitude (Whitman et al., 1998; Curtis et al., 2006).”
BackgroundSharma, M., Mitnala, S., Vishnubhotla, R., Mukherjee, R., Reddy, D. N., & Rao, P. N. (2015) · Journal of Clinical and Experimental Hepatology · doi:10.1016/j.jceh.2015.02.002 · Cited by 154
“The intestinal microbiome is composed of bacteria, viruses, yeast and archae.(144,145) Recently, there has been a growing body of evidence that implicate gut microbiota in NAFLD and its progression to NASH.”
BackgroundBiogenic methane in shale gas and coal bed methane: A review of current knowledge and gaps
Colosimo, F., Thomas, R., Lloyd, J. R., Taylor, K. G., Boothman, C., Smith, A. D., Lord, R., & Kalin, R. M. (2016) · International Journal of Coal Geology · doi:10.1016/j.coal.2016.08.011 · Cited by 156
“It has been suggested that the biomass into the deep biosphere constitutes one-tenth (Parkes et al., 2000), or even one-third (Whitman et al., 1998) of the total global, living biomass.”
BackgroundOrellana, L. H., Chee‐Sanford, J. C., Sanford, R. A., Löffler, F. E., & Konstantinidis, K. T. (2017) · Applied and Environmental Microbiology · doi:10.1128/aem.01646-17 · Cited by 148
“soil ecosystems has been extensively reported (3, 4), the dynamics of microbial communities responding to seasonal agricultural perturbations (e.”
BackgroundGlacier ice archives nearly 15,000-year-old microbes and phages
Zhong, Z.-P., Tian, F., Roux, S., Gazitúa, M. C., Solonenko, N., Li, Y.-F., Davis, M. E., Etten, J. L. V., Mosley‐Thompson, E., Rich, V. I., et al. (2021) · Microbiome · doi:10.1186/s40168-021-01106-w · Cited by 157
“These studies revealed microbial cell concentrations of 10(2) to 10(4) cells ml in most glacier-ice samples [4], which are several orders of magnitude lower than other environments such as seawater or soils [7].”
BackgroundDeep subseafloor microbial cells on physiological standby
Jørgensen, B. B. (2011) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1115421108 · Cited by 141
“M icroscopic counts of microbial cells in deep sediment cores retrieved by scientific ocean drilling have revealed the largest living ecosystem on earth: the deep biosphere (1, 2).”
BackgroundBrown, E. M., McTaggart, L. R., Zhang, S. X., Low, D. E., Stevens, D. A., & Richardson, S. E. (2013) · PLoS ONE · doi:10.1371/journal.pone.0059237 · Cited by 148
The human microbiome and bile acid metabolism: Dysbiosis, dysmetabolism, disease and intervention
Jones, M. L., Martoni, C. J., Ganopolsky, J. G., Labbé, A., & Prakash, S. (2014) · Expert Opinion on Biological Therapy · doi:10.1517/14712598.2014.880420 · Cited by 150
“All surfaces of the human body that are exposed to the environment are colonized, including skin, respiratory system, urogenital tract and gastrointestinal (GI) tract, totaling at least 100 trillion microbial cells [6].”
BackgroundA physiological perspective on the origin and evolution of photosynthesis
Martin, W. F., Bryant, D. A., & Beatty, J. T. (2017) · FEMS Microbiology Reviews · doi:10.1093/femsre/fux056 · Cited by 158
Einen, J., Thorseth, I. H., & Øvreås, L. (2008) · FEMS Microbiology Letters · doi:10.1111/j.1574-6968.2008.01119.x · Cited by 147
The Genomic Enzymology of Antibiotic Resistance
Morar, M., & Wright, G. D. (2010) · Annual Review of Genetics · doi:10.1146/annurev-genet-102209-163517 · Cited by 159
Milici, M., Deng, Z.-L., Tomasch, J., Decelle, J., Wos‐Oxley, M. L., Wang, H., Jáuregui, R., Plumeier, I., Giebel, H., Badewien, T. H., et al. (2016) · Frontiers in Microbiology · doi:10.3389/fmicb.2016.00649 · Cited by 146
“In the ocean, prokaryotes (bacteria and archaea) are the most abundant living organisms, with an average of 5 × 10 5 cells per milliliter of sea water on the continental shelf but also in the upper 200 m of the oceanic water column (Whitman et al., 1998).”
BackgroundMicrobial community diversity in seafloor basalt from the Arctic spreading ridges
Lysnes, K., Thorseth, I. H., Steinsbu, B. O., Øvreås, L., Torsvik, T., & Pedersen, R. B. (2004) · FEMS Microbiology Ecology · doi:10.1016/j.femsec.2004.06.014 · Cited by 155
“stutzeri are common constituents of laboratory water supply [61–63] and have repeatedly been found to contaminate low-biomass samples [2,61].”
BackgroundA Starving Majority Deep Beneath the Seafloor
Jørgensen, B. B., & D'Hondt, S. L. (2006) · Science · doi:10.1126/science.1133796 · Cited by 146
Deep-Sea Hydrothermal Vents: Potential Hot Spots for Natural Products Discovery?
Thornburg, C. C., Zabriskie, T. M., & McPhail, K. L. (2010) · Journal of Natural Products · doi:10.1021/np900662k · Cited by 149
Soil Microbial Community Responses to a Decade of Warming as Revealed by Comparative Metagenomics
Luo, C., Rodriguez‐R, L. M., Johnston, E. R., Wu, L., Cheng, L., Xue, K., Tu, Q., Deng, Y., He, Z., Shi, Z. J., et al. (2013) · Applied and Environmental Microbiology · doi:10.1128/aem.03712-13 · Cited by 153
Carbonate Mineral Precipitation for Soil Improvement Through Microbial Denitrification
Hamdan, N., Kavazanjian, E., Rittmann, B. E., & Karatas, I. (2016) · Geomicrobiology Journal · doi:10.1080/01490451.2016.1154117 · Cited by 147
“Estimates suggest that nearly half of the Earth’s biomass is comprised of microorganisms found in the subsurface and oceanic subsurface (Whitman et al. 1998).”
BackgroundChemical Challenges to Bacterial AHL Signaling in the Environment
Decho, A. W., Frey, R. L., & Ferry, J. L. (2010) · Chemical Reviews · doi:10.1021/cr100311q · Cited by 146
Hu, P., Tom, L., Singh, A., Thomas, B., Baker, B., Piceno, Y., Andersen, G., & Banfield, J. (2016) · mBio · doi:10.1128/mbio.01669-15 · Cited by 135
Microbial competition in porous environments can select against rapid biofilm growth
Coyte, K. Z., Tabuteau, H., Gaffney, E. A., Foster, K. R., & Durham, W. M. (2016) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1525228113 · Cited by 146
“The overwhelming majority of bacteria live in porous environments between the particles that compose soil, aquifers, and sediments, and cumulatively comprise roughly half of the carbon within living organisms globally (8).”
BackgroundEnergy and carbon metabolisms in a deep terrestrial subsurface fluid microbial community
Momper, L., Jungbluth, S. P., Lee, M., & Amend, J. P. (2017) · The ISME Journal · doi:10.1038/ismej.2017.94 · Cited by 143
“Early estimates of terrestrial subsurface cells were on the order of 0.25–2.5 × 1030 (Whitman et al., 1998).”
BackgroundInfluentialMetagenomics of extreme environments
Cowan, D. A., Ramond, J., Makhalanyane, T. P., & Maayer, P. D. (2015) · Current Opinion in Microbiology · doi:10.1016/j.mib.2015.05.005 · Cited by 148
Airborne desert dust and aeromicrobiology over the Turkish Mediterranean coastline
Griffin, D. W., Kubilay, N., Koçak, M., Gray, M., Borden, T. C., & Shinn, E. A. (2007) · Atmospheric Environment · doi:10.1016/j.atmosenv.2007.01.023 · Cited by 147
“At estimates of 10(9) bacteria and 10(6) fungi per gram of top soil it should not be surprising that when soil is mobilized into the atmosphere by storm activity, concentrations of airborne microorganisms increase in the environment and later in the downwind regions (Tate, 2000; Whitman et al., 1998).”
A metagenomic window into carbon metabolism at 3 km depth in Precambrian continental crust
Magnabosco, C., Ryan, K., Lau, M. C. Y., Kuloyo, O., Lollar, B. S., Kieft, T. L., van Heerden, E., & Onstott, T. C. (2015) · The ISME Journal · doi:10.1038/ismej.2015.150 · Cited by 148
“…the habitable, yet carbon poor, regions of the deep continental crust and the previous estimates of subsurface microbial turnover times (1–2 × 103 years; Whitman et al., 1998), the deep continental subsurface microbial carbon metabolism requires reevaluation from an interdisciplinary perspective.”
Background
Metadata
- Authors
- Whitman, W. B., Coleman, D. C., & Wiebe, W. J.
- Affiliations
- University of Georgia (US)
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 1998-06-09
- Volume
- 95
- Issue
- 12
- Pages
- 6578-6583
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC33863
- OpenAlex
- W2118001436
- Semantic Scholar
- 321e4c5e6031364c39db3cfdb151dcc2deaac1ba
- Cited by
- 4,871
- References
- 92
- Citations by year
- 2026: 86 · 2025: 134 · 2024: 159 · 2023: 220 · 2022: 190 · 2021: 252 · 2020: 223 · 2019: 238 · 2018: 273 · 2017: 251 · 2016: 281 · 2015: 282 · 2014: 311 · 2013: 252 · 2012: 238
- FWCI
- 6.26
- Type
- journal article
- Language
- en
- Open access
- green
- europepmc.org
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Physical Sciences › Environmental Science › Ecology › Microbial Community Ecology and Physiology
- Life Sciences › Agricultural and Biological Sciences › Soil Science › Soil Carbon and Nitrogen Dynamics
- Physical Sciences › Earth and Planetary Sciences › Oceanography › Marine Plankton and Biogeochemistry
- Keywords
- prokaryotes, nitrogen, phosphorus, open ocean, soil, subsurface environments, genetic diversity
- MeSH terms
- Prokaryotic Cells
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