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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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“As a group, bacteria are the most genetically diverse and abundant life form on Earth [1].”
BackgroundThe Importance of Chitin in the Marine Environment
Souza, C. P., de Almeida, B. C., Colwell, R. R., & Rivera, I. N. G. (2011) · Marine Biotechnology · doi:10.1007/s10126-011-9388-1 · Cited by 227
“Thus, this environment constitutes an inestimable source for the discovery of new microorganisms and bioactive compounds for biotechnology and sustainable development (Colwell 1997; Whitman et al. 1998 ).”
BackgroundInfluentialSoil biodiversity: functions, threats and tools for policy makers
Turbé, A., Toni, A. D., Benito, P., Lavelle, P., Lavelle, P., Ruiz, N., van der Putten, W. H., Labouze, É., & Mudgal, S. (2010) · HAL · Cited by 221
Modeling Taxa-Abundance Distributions in Microbial Communities using Environmental Sequence Data
Sloan, W. T., Woodcock, S., Lunn, M., et al. (2006) · Microbial Ecology · doi:10.1007/s00248-006-9141-x · Cited by 220
“The computational expense [19] of this discrete MarkovChain formulation makes it impossible to apply to the very large diverse populations that typify the microbial world [27].”
BackgroundStructure, Function, and Biology of the Enterococcus faecalis Cytolysin
Tyne, D. V., Martin, M. J., & Gilmore, M. S. (2013) · Toxins · doi:10.3390/toxins5050895 · Cited by 236
“colonized with up to 10(11) bacterial cells/gram feces [3,4].”
BackgroundThe Altered Schaedler Flora: Continued Applications of a Defined Murine Microbial Community
Brand, M. W., Wannemuehler, M. J., Phillips, G. J., Proctor, A., Overstreet, A.-M. C., Jergens, A. E., Orcutt, R. P., & Fox, J. G. (2015) · ILAR Journal · doi:10.1093/ilar/ilv012 · Cited by 223
Marine bacteria exhibit a bipolar distribution
Sul, W. J., Oliver, T. A., Ducklow, H. W., Amaral‐Zettler, L., & Sogin, M. L. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1212424110 · Cited by 218
“Arguments for “everything is everywhere” cite enormousmicrobial population sizes (10) or the presence of strong passive dispersal forces (i.”
BackgroundComparative genomics and evolution of the tailed-bacteriophages
Casjens, S. (2005) · Current Opinion in Microbiology · doi:10.1016/j.mib.2005.06.014 · Cited by 218
Extraordinary phylogenetic diversity and metabolic versatility in aquifer sediment
Castelle, C., Hug, L., Wrighton, K., Thomas, B., Williams, K., Wu, D., Tringe, S., Singer, S., Eisen, J., & Banfield, J. (2013) · Nature Communications · doi:10.1038/ncomms3120 · Cited by 224
Global distribution of nearly identical phage-encoded DNA sequences
Breitbart, M., Miyake, J. H., & Rohwer, F. L. (2004) · FEMS Microbiology Letters · doi:10.1111/j.1574-6968.2004.tb09654.x · Cited by 217
“There are an estimated 10(30) prokaryotes in the biosphere, the vast majority of which are in the subsurface terrestrial, sediments, and the oceans [18].”
BackgroundFrom peds to paradoxes: Linkages between soil biota and their influences on ecological processes
Coleman, D. C. (2007) · Soil Biology and Biochemistry · doi:10.1016/j.soilbio.2007.08.005 · Cited by 212
“5, ARTICLE IN PRESS D.C. Coleman / Soil Biology & Biochemistry 40 (2008) 271–289272 Whitman et al., 1998)?”
BackgroundSelf-citationCarbon and nitrogen assimilation in deep subseafloor microbial cells
Morono, Y., Terada, T., Nishizawa, M., Ito, M., Hillion, F., Takahata, N., Sano, Y., & Inagaki, F. (2011) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1107763108 · Cited by 218
Zhou, J., Lyu, Y., Richlen, M. L., Anderson, D. M., & Cai, Z. (2016) · Critical Reviews in Plant Sciences · doi:10.1080/07352689.2016.1172461 · Cited by 217
“0 £ 10(30) cells, which accounts for approximately half of the total biomass of living organisms on the Earth (Whitman et al., 1998).”
BackgroundInfluentialRigsbee, L., Agans, R., Shankar, V., Kenche, H., Khamis, H. J., Michail, S., & Paliy, O. (2012) · American Journal of Gastroenterology · doi:10.1038/ajg.2012.287 · Cited by 207
Commensal bacteria mediated defenses against pathogens
Abt, M. C., & Pamer, E. G. (2014) · Current Opinion in Immunology · doi:10.1016/j.coi.2014.03.003 · Cited by 223
“The mammalian gastrointestinal tract harbors one of the densest microbial ecosystems found in nature, peaking in the proximal colon at 10(11)–10(12) cells/mL [1].”
BackgroundFriend Turned Foe: Evolution of Enterococcal Virulence and Antibiotic Resistance
Tyne, D. V., & Gilmore, M. S. (2014) · Annual Review of Microbiology · doi:10.1146/annurev-micro-091213-113003 · Cited by 223
“As a typical omnivorous mammal, humans shed the equivalent of their entire intestinal microbiota into the environment every 2–3 weeks (37, 130).”
BackgroundBiological residues define the ice nucleation properties of soil dust
Conen, F., Morris, C. E., Leifeld, J., Yakutin, M. V., & Alewell, C. (2011) · Atmospheric chemistry and physics · doi:10.5194/acp-11-9643-2011 · Cited by 213
Expression and isolation of antimicrobial small molecules from soil DNA libraries.
MacNeil, I. A., Tiong, C. L., Minor, C., August, P. R., Grossman, T. H., Loiacono, K. A., Lynch, B. A., Phillips, T., Narula, S. S., Sundaramoorthi, R., et al. (2001) · PubMed · Cited by 212
The rational exploration of microbial diversity
Quince, C., Curtis, T. P., & Sloan, W. T. (2008) · The ISME Journal · doi:10.1038/ismej.2008.69 · Cited by 212
“The microbial world is vast, with 10 organisms present on the Earth (Whitman et al., 1998), diverse (Curtis et al.”
BackgroundSampling the mobile gene pool: innovation via horizontal gene transfer in bacteria
Hall, J. P. J., Brockhurst, M. A., & Harrison, E. (2017) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2016.0424 · Cited by 224
Culturing captures members of the soil rare biosphere
Shade, A. L., Hogan, C. S., Klimowicz, A. K., Linske, M., McManus, P. S., & Handelsman, J. (2012) · Environmental Microbiology · doi:10.1111/j.1462-2920.2012.02817.x · Cited by 211
“Microorganisms are the most abundant organisms on Earth and represent an unfathomably high level of diversity (Whitman et al., 1998; Rappé and Giovannoni, 2003; Schloss and Handelsman, 2007).”
BackgroundBrazelton, W. J., Nelson, B., & Schrenk, M. O. (2012) · Frontiers in Microbiology · doi:10.3389/fmicb.2011.00268 · Cited by 219
“The potentially vast microbial diversity and biomass of the subsurface biosphere (Whitman et al., 1998) has been frequently noted (Biddle et al., 2006; Huber et al., 2007; Santelli et al., 2008; Schrenk et al., 2010), but there is very little evidence to indicate how much of it is supported by new…”
BackgroundDombrowski, N., Seitz, K. W., Teske, A., & Baker, B. J. (2017) · Microbiome · doi:10.1186/s40168-017-0322-2 · Cited by 211
Modulation of the human gut microbiota by phenolics and phenolic fiber‐rich foods
Loo, Y. T., Howell, K. S., Chan, M., Zhang, P., & Ng, K. (2020) · Comprehensive Reviews in Food Science and Food Safety · doi:10.1111/1541-4337.12563 · Cited by 223
Uterine Microbiota Progression from Calving until Establishment of Metritis in Dairy Cows
Jeon, S. J., Vieira-Neto, A., Gobikrushanth, M., et al. (2015) · Applied and Environmental Microbiology · doi:10.1128/aem.01753-15 · Cited by 209
“Although culture-based studies have laid out the foundation of our understanding of the uterine microbiota, previous studies might have underestimated the microbial complexity of the intrauterine environment of cows postpartum, given that less than 1% of the microorganisms in many environments are readily cultured und…”
Konstantinidis, K. T., Braff, J. C., Karl, D. M., & DeLong, E. F. (2009) · Applied and Environmental Microbiology · doi:10.1128/aem.00473-09 · Cited by 215
“Microbes dominate the oceans’ interior (42) and thus drive the biochemical cycles that sustain life in this habitat.”
BackgroundEnvironmental distribution of prokaryotic taxa
Tamames, J., Abellan, J. J., Pignatelli, M., Camacho, A., & Moyá, A. (2010) · BMC Microbiology · doi:10.1186/1471-2180-10-85 · Cited by 208
“Also the massive number of existing species makes cataloguing microbial diversity difficult [18].”
BackgroundBiological control in the microbiome era: Challenges and opportunities
Massart, S., Martínez-Medina, M., & Jijakli, M. H. (2015) · Biological Control · doi:10.1016/j.biocontrol.2015.06.003 · Cited by 218
“the phyllosphere (Lindow and Brandl, 2003) and 10(6)–10(9) cells/g in the rhizosphere (Whitman et al., 1998).”
BackgroundA Comparison of Methods for Clustering 16S rRNA Sequences into OTUs
Chen, W., Zhang, H., Cheng, Y., Zhang, S., & Zhao, H. (2013) · PLoS ONE · doi:10.1371/journal.pone.0070837 · Cited by 207
“earth and more diverse than any other organisms [1].”
BackgroundHalophilic Bacteria as a Source of Novel Hydrolytic Enzymes
Moreno, M. D. L., Pérez, D., Garcia, M. B., et al. (2013) · Life · doi:10.3390/life3010038 · Cited by 206
Yang, Y., Wang, N., Guo, X., et al. (2017) · PLoS ONE · doi:10.1371/journal.pone.0178425 · Cited by 206
Microbial Growth under Limiting Conditions-Future Perspectives
Grau, J. M. G., & Aranda, B. (2023) · Microorganisms · doi:10.3390/microorganisms11071641 · Cited by 215
“Growth is a result of microbial activity [1,26], but quantification of the growth status of a microbial population or an individual cell is an issue that requires further investigation and remains a major challenge to be solved.”
BackgroundComparison of microbial numbers in soils by using various culture media and temperatures
Vieira, F. C. S., & Nahas, E. (2005) · Microbiological Research · doi:10.1016/j.micres.2005.01.004 · Cited by 205
“Bacterial counts in different soils ranged from 4 106 to 2 109 g 1 dry soil (Whitman et al., 1998).”
BackgroundIsolation and characterization of low nucleic acid (LNA)-content bacteria
Wang, Y., Hammes, F., Boon, N., Chami, M., & Egli, T. W. (2009) · The ISME Journal · doi:10.1038/ismej.2009.46 · Cited by 204
“Planktonic bacteria are ubiquitous in aquatic environments and an average density of 10(6)–10(7) cells ml is typically observed in surface freshwater (Whitman et al., 1998; Wang et al., 2007).”
BackgroundThe human gut virome: a multifaceted majority
Ogilvie, L. A., & Jones, B. V. (2015) · Frontiers in Microbiology · doi:10.3389/fmicb.2015.00918 · Cited by 212
“This viral community also appears to encode a surprisingly rich functional repertoire that confers a range of attributes to their bacterial hosts, ranging from bacterial virulence and pathogenesis to maintaining host–microbiome stability and community resilience.”
BackgroundMeasuring the biodiversity of microbial communities by flow cytometry
Props, R., Monsieurs, P., Mysara, M., Clement, L., & Boon, N. (2016) · Methods in Ecology and Evolution · doi:10.1111/2041-210x.12607 · Cited by 203
Microbial oceanography: paradigms, processes and promise
Karl, D. M. (2007) · Nature Reviews Microbiology · doi:10.1038/nrmicro1749 · Cited by 203
“Ocean habitats and their microbial assemblages Microorganisms inhabit all marine ecosystems, from the tropics to the sea ice and from the well-lit surface waters to the deep abyss; they truly are the “unseen majority&rdquo”
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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