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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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Prokaryotic responses to hydrostatic pressure in the ocean – a review
Tamburini, C., Boutrif, M., Garel, M., Colwell, R. R., & Deming, J. W. (2013) · Environmental Microbiology · doi:10.1111/1462-2920.12084 · Cited by 183
“3 ¥ 10 m) (Whitman et al., 1998).”
InfluentialSantos, T. M. A., Gilbert, R. O., & Bicalho, R. C. (2010) · Journal of Dairy Science · doi:10.3168/jds.2010-3668 · Cited by 183
“…culturing, previous studies might have underestimated the microbial complexity of the intrauterine environment of postpartum cows, given that less than 1% of the microorganisms in many environments are readily cultured under standard laboratory conditions (Amann et al., 1995; Whitman et al., 1998).”
BackgroundThe source and transport of bioaerosols in the air: A review
Xie, W., Li, Y., Bai, W., Hou, J., Ma, T., Zeng, X., Zhang, L., & An, T. (2020) · Frontiers of Environmental Science & Engineering · doi:10.1007/s11783-020-1336-8 · Cited by 187
Microbial biofilms for electricity generation from water evaporation and power to wearables
Liu, X., Ueki, T., Gao, H., Woodard, T. L., Nevin, K. P., Fu, T., Fu, S., Sun, L., Lovley, D. R., & Yao, J. (2022) · Nature Communications · doi:10.1038/s41467-022-32105-6 · Cited by 183
Combined constraints on global ocean primary production using observations and models
Buitenhuis, E. T., Hashioka, T., & Quéré, C. L. (2013) · Global Biogeochemical Cycles · doi:10.1002/gbc.20074 · Cited by 191
Cyanobacteria as Biocatalysts for Carbonate Mineralization
Kamennaya, N. A., Ajo-Franklin, C. M., Northen, T. R., & Jansson, C. (2012) · Minerals · doi:10.3390/min2040338 · Cited by 191
“It is estimated that half of global photosynthesis is carried out by phytoplankton [46], with cyanobacteria being, at times, the dominant contributors to”
BackgroundNew techniques in sediment core analysis: an introduction
Rothwell, R. G., & Rack, F. R. (2006) · Geological Society London Special Publications · doi:10.1144/gsl.sp.2006.267.01.01 · Cited by 182
Grau, J. M. G., Portillo, M. C., Belda‐Ferre, P., et al. (2012) · PLoS ONE · doi:10.1371/journal.pone.0029973 · Cited by 181
“Microorganisms play a major role in the functioning of biochemical cycles of elements [1] involving a large variety of microbial taxa.”
BackgroundAvershina, E., Storrø, O., Øien, T., Johnsen, R., Pope, P. B., & Rudi, K. (2013) · FEMS Microbiology Ecology · doi:10.1111/1574-6941.12223 · Cited by 192
“While it is widely accepted that the human gut is one of the most densely populated bacterial communities on the Earth (Whitman et al., 1998), the general mechanisms for host–bacterial interactions are not yet completely described (Avershina & Rudi, 2013).”
BackgroundBifunctional antimicrobial conjugates and hybrid antimicrobials
Klahn, P., & Brönstrup, M. (2017) · Natural Product Reports · doi:10.1039/c7np00006e · Cited by 183
Macroecological laws describe variation and diversity in microbial communities
Grilli, J. (2020) · Nature Communications · doi:10.1038/s41467-020-18529-y · Cited by 170
Bacterial-nanostructure interactions: The role of cell elasticity and adhesion forces
Elbourne, A. J., Chapman, J., Gelmi, A. A., Cozzolino, D., Crawford, R. J., & Truong, V. K. (2019) · Journal of Colloid and Interface Science · doi:10.1016/j.jcis.2019.03.050 · Cited by 184
Orr, C. H., James, A. F., Leifert, C., Cooper, J. M., & Cummings, S. P. (2010) · Applied and Environmental Microbiology · doi:10.1128/aem.01250-10 · Cited by 189
“16S rRNA copy numbers are also are lower than would be expected in agricultural soil and are more similar to numbers observed in forest soil (49).”
BackgroundHomeland Security: IgA immunity at the frontiers of the body
Macpherson, A. J., Geuking, M. B., & McCoy, K. D. (2012) · Trends in Immunology · doi:10.1016/j.it.2012.02.002 · Cited by 174
Gastrointestinal Tract Microbiota and Probiotics in Production Animals
Yeoman, C. J., & White, B. A. (2013) · Annual Review of Animal Biosciences · doi:10.1146/annurev-animal-022513-114149 · Cited by 193
Redox Fluctuations Control the Coupled Cycling of Iron and Carbon in Tropical Forest Soils
Bhattacharyya, A., Campbell, A., Tfaily, M., Lin, Y., Kukkadapu, R., Silver, W. L., Nico, P., & Pett‐Ridge, J. (2018) · Environmental Science & Technology · doi:10.1021/acs.est.8b03408 · Cited by 183
Webster, G., Embley, T. M., Freitag, T. E., et al. (2005) · Environmental Microbiology · doi:10.1111/j.1462-2920.2005.00740.x · Cited by 178
“Realization and acceptance of the importance of prokaryotes in global biogeochemical cycling (Whitman et al ., 1998) has coincided with the development of molecular techniques that have demonstrated considerable prokaryotic diversity and the prevalence of uncultured groups (Pace, 1997; Hugenholtz…”
BackgroundBakker, P. A. H. M., Ran, L., Pieterse, C. M. J., et al. (2003) · Canadian Journal of Plant Pathology · doi:10.1080/07060660309507043 · Cited by 178
Layer-by-layer assemblies for antibacterial applications
Zhu, X., & Loh, X. J. (2015) · Biomaterials Science · doi:10.1039/c5bm00307e · Cited by 181
Benkendorff, K. (2010) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/j.1469-185x.2010.00124.x · Cited by 176
Intestinal M cells: The fallible sentinels?
Miller, H. (2007) · World Journal of Gastroenterology · doi:10.3748/wjg.v13.i10.1477 · Cited by 175
Shen, Z., Penton, C. R., Lv, N., Xue, C., Yuan, X., Ruan, Y., Li, R., & Shen, Q. (2017) · Microbial Ecology · doi:10.1007/s00248-017-1052-5 · Cited by 175
Review of ground improvement using microbial induced carbonate precipitation (MICP)
Wang, Z., Zhang, N., Cai, G., Jin, Y. J., Ding, N., & Shen, D. (2017) · Marine Georesources and Geotechnology · doi:10.1080/1064119x.2017.1297877 · Cited by 176
“The number is reduced from 1011 to 106 at depth 2–30 m (Whitman, Coleman, and Weibe 1998; Mitchell and Santamarina 2005).”
BackgroundReinthaler, T., van Aken, H., Veth, C., Arı́stegui, J., Robinson, C., Williams, P., Lebaron, P., & Herndl, G. J. (2006) · Limnology and Oceanography · doi:10.4319/lo.2006.51.3.1262 · Cited by 174
Small but Mighty: Cell Size and BacteriaPreprint
Levin, P. A., & Angert, E. R. (2015) · Cold Spring Harbor Perspectives in Biology · doi:10.1101/cshperspect.a019216 · Cited by 176
“Despite their astounding global abundance ( 1030 cells) and their substantial contribution to the total biomass of planet earth (Whitman et al. 1998; Kallmeyer et al. 2012), our inability to see these tiny life forms shrouded their nearly limitless diversity in mystery.”
BackgroundChyba, C. F., & Phillips, C. B. (2002) · Origins of Life and Evolution of Biospheres · doi:10.1023/a:1013958519734 · Cited by 171
“Integrated over the history of the Solar System, some 1012 kg of carbon should have been accumulated by Europa in this way (Pierazzo and Chyba, 2001) – about 0.1% of the carbon present in Earth’s total biomass(Whitman et al., 1998).”
BackgroundMethodInfluentialMolecular Bases and Role of Viruses in the Human Microbiome
Abeles, S. R., & Pride, D. T. (2014) · Journal of Molecular Biology · doi:10.1016/j.jmb.2014.07.002 · Cited by 171
“There are an estimated 10(31) phage on earth, based on calculations of 10(30) bacteria on the planet [15] and approximately 10 phage that exist for every bacteria [16].”
BackgroundSize doesn’t matter: towards a more inclusive philosophy of biology
O’Malley, M. A., & Dupré, J. (2007) · Biology & Philosophy · doi:10.1007/s10539-006-9031-0 · Cited by 176
“(8)A commonly accepted estimate is 4–6· 10(30) prokaryote cells in all habitats (Whitman et al. 1998) and 4· 10(30) viruses just in ocean waters (Suttle 2005).”
BackgroundAmachi, S. (2008) · Microbes and Environments · doi:10.1264/jsme2.me08548 · Cited by 174
Microbial ecology of the atmosphere
Šantl‐Temkiv, T., Amato, P., Casamayor, E. O., Lee, P. K. H., & Pointing, S. B. (2022) · FEMS Microbiology Reviews · doi:10.1093/femsre/fuac009 · Cited by 190
“Nonetheless, a global scale meta-analysis of bacterial abundance across biomes indicated ∼5 × 1019 cells may occur globally in the atmospheric boundary layer, compared with ∼1.2 × 1029 in oceans and 2.6 × 1029 in soils globally (Whitman et al. 1998).”
BackgroundInfluentialMicrobial interactions within beneficial consortia promote soil health
Wu, D., Wang, W., Yao, Y., Li, H., Wang, Q., & Niu, B. (2023) · Science of The Total Environment · doi:10.1016/j.scitotenv.2023.165801 · Cited by 181
Phage Community Dynamics in Hot Springs
Breitbart, M., Wegley, L., Leeds, S. G., et al. (2004) · Applied and Environmental Microbiology · doi:10.1128/aem.70.3.1633-1640.2004 · Cited by 173
“These rates are similar to those estimated by a completely independent approach based on data from groundwater (52).”
ResultHundreds of genes experienced convergent shifts in selective pressure in marine mammals
Chikina, M., Robinson, J. D., & Clark, N. L. (2016) · Molecular Biology and Evolution · doi:10.1093/molbev/msw112 · Cited by 171
“Indeed, the oceans harbor an enormous diversity and number of viruses and prokaryotes, which could frequently come in to contact with marine mammal skin (Whitman et al. 1998; Suttle 2007).”
BackgroundAsexual reproduction: Genetics and evolutionary aspects
Meeûs, T. D., Prugnolle, F., & Agnew, P. (2007) · Cellular and Molecular Life Sciences · doi:10.1007/s00018-007-6515-2 · Cited by 184
Sun, Y., Cai, Y., Huse, S. M., Knight, R. D., Farmerie, W. G., Wang, X., & Mai, V. (2011) · Briefings in Bioinformatics · doi:10.1093/bib/bbr009 · Cited by 172
Abd‐Alla, M. H., Al-Amri, S. M., & El-Enany, A. E. (2023) · Agriculture · doi:10.3390/agriculture13112092 · Cited by 205
“They include heterotrophic bacteria, such as Azotobacter , and autotrophic cyanobacteria Heterotrophic bacteria obtain energy from organic compounds, while cyanobacteria can utilize sunlight as an energy source [43,44]; 2.”
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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