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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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Männik, J., Driessen, R. P. C., Galajda, P., Keymer, J. E., & Dekker, C. (2009) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.0907542106 · Cited by 300
“These bacteria constitute a large portion of the Earth’s biomass (2) and are essential for the functioning of soil.”
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“of human health and the environment [1, 2].”
Phasing amplicon sequencing on Illumina Miseq for robust environmental microbial community analysis
Wu, L., Wen, C., Qin, Y., Yin, H., Tu, Q., Nostrand, J. D. V., Yuan, T., Yuan, M., Deng, Y., & Zhou, J. Z. (2015) · BMC Microbiology · doi:10.1186/s12866-015-0450-4 · Cited by 298
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Lechtenfeld, O. J., Hertkorn, N., Shen, Y., Witt, M., & Benner, R. (2015) · Nature Communications · doi:10.1038/ncomms7711 · Cited by 291
Metagenomic gene discovery: past, present and future
Cowan, D. A., Meyer, Q. C., Stafford, W. H. L., Samson, M., Cameron, R. A., & Wittwer, P. (2005) · Trends in biotechnology · doi:10.1016/j.tibtech.2005.04.001 · Cited by 289
“The total number of prokaryotic cells on earth has been estimated at 4–6!1030 [ 1 ], thought to comprise between 106 and 108 separate genospecies (distinct taxonomic”
BackgroundExpanding the soil antibiotic resistome: exploring environmental diversity
D’Costa, V. M., Griffiths, E., & Wright, G. D. (2007) · Current Opinion in Microbiology · doi:10.1016/j.mib.2007.08.009 · Cited by 290
“Antibiotic producing bacteria: a reservoir and putative origin of resistance determinants Inhabited by up to 10(9) microorganisms/g [9], the diversity of microbial life concealed within the soil has been explored in the search for new clinical and medicinal applications.”
BackgroundEcology and Physics of Bacterial Chemotaxis in the Ocean
Stocker, R., & Seymour, J. R. (2012) · Microbiology and Molecular Biology Reviews · doi:10.1128/mmbr.00029-12 · Cited by 293
Earle, S. G., Wu, C., Charlesworth, J., Stoesser, N. E., Gordon, N. C., Walker, T. M., Spencer, C. C. A., Iqbal, Z., Clifton, D. A., Hopkins, K. L., et al. (2016) · Nature Microbiology · doi:10.1038/nmicrobiol.2016.41 · Cited by 287
“encoded as a vector indicating the presence (1) or absence (0) of the first (K−1) alleles and βl becomes a vector of length (K−1).”
Critical review: Microbially influenced corrosion of buried carbon steel pipes
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Energy for microbial life on Europa
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Gene expression in the deep biosphere
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Ecology of viruses in soils: Past, present and future perspectives
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“2 × 10 tC in the open ocean are estimated (Whitman et al. 1998).”
BackgroundInfluentialElevated Abundance of Bacteriophage Infecting Bacteria in Soil
Ashelford, K. E., Day, M. J., & Fry, J. C. (2003) · Applied and Environmental Microbiology · doi:10.1128/aem.69.1.285-289.2003 · Cited by 281
“In view of the large biomass of bacteria in the biosphere (30), our results indicate that predation of bacteria by viruses will be an important factor in controlling and stimulating the growth of bacterial populations in soil.”
BackgroundMethanogenic pathways of coal-bed gas in the Powder River Basin, United States: The geologic factor
Flores, R. M., Rice, C. A., Stricker, G. D., et al. (2008) · International Journal of Coal Geology · doi:10.1016/j.coal.2008.02.005 · Cited by 278
Unlocking the potential of metagenomics through replicated experimental design
Knight, R. D., Jansson, J. K., Field, D., Fierer, N., Desai, N., Fuhrman, J. A., Hugenholtz, P., van der Lelie, D., Meyer, F., Stevens, R. L., et al. (2012) · Nature Biotechnology · doi:10.1038/nbt.2235 · Cited by 294
Groundwater activity on Mars and implications for a deep biosphere
Michalski, J. R., Cuadros, J., Niles, P. B., Parnell, J., Rogers, A. D., & WRIGHT, S. P. (2013) · Nature Geoscience · doi:10.1038/ngeo1706 · Cited by 298
Rickettsial evolution in the light of comparative genomics
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Weighing the deep continental biosphere
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“Using data from a single sandstone aquifer (Hazen et al., 1991), Whitman et al. (1998) determined that 0.058% of cells in Table 2.”
BackgroundMethodInfluentialThe Role of Gut Microbiota on Insulin Resistance
Caricilli, A. M., & Saad, M. A. (2013) · Nutrients · doi:10.3390/nu5030829 · Cited by 285
“Thus, a new approach was introduced, culture-independent sequencing [21–23], which made detection of microbial genes and disease-associated patterns in our gut microbiota possible.”
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“Bacteria are the most diverse organisms on the planet [1].”
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Jørgensen, B. B. (2006) · Springer eBooks · doi:10.1007/3-540-32144-6_5 · Cited by 279
“Yet, calculations from cell numbers and measured or modeled mineralization rates show that the organic carbon flux available in million year old sediments allows only generation times of years to thousands of years ( Whitman et al. 1998; Schippers et al. 2005).”
BackgroundCultivating the uncultured: limits, advances and future challenges
Alain, K., & Quérellou, J. (2009) · Extremophiles · doi:10.1007/s00792-009-0261-3 · Cited by 279
“…have been described (and validated by the International Committee on Systematics of Prokaryotes) in the one hundred and twenty years, since the invention of the Petri dish (Achtman & Wagner, 2008), even though we now realize that prokaryotes represent “the unseen majority” (Whitman et al., 1998).”
BackgroundBacteriophage and their lysins for elimination of infectious bacteria
O’Flaherty, S., Ross, R. P., & Coffey, A. (2009) · FEMS Microbiology Reviews · doi:10.1111/j.1574-6976.2009.00176.x · Cited by 272
A census-based estimate of Earth's bacterial and archaeal diversity
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Finding alternatives to antibiotics
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Subramani, R., & Sipkema, D. (2019) · Marine Drugs · doi:10.3390/md17050249 · Cited by 299
Accounting for reciprocal host–microbiome interactions in experimental science
Stappenbeck, T. S., & Virgin, H. W. (2016) · Nature · doi:10.1038/nature18285 · Cited by 272
PHACTS, a computational approach to classifying the lifestyle of phages
McNair, K. A., Bailey, B., & Edwards, R. A. (2012) · Bioinformatics · doi:10.1093/bioinformatics/bts014 · Cited by 269
“[13:23 25/2/2012 Bioinformatics-bts014.tex] Page: 614 614–618 Motivation: Bacteriophages have two distinct lifestyles: virulent and temperate.”
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“S y st e m [4 3 ,5 3 – 5 5 ] E a st e rn E q u a to ri a l”
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BUTTERFIELD, N. J. (2007) · Palaeontology · doi:10.1111/j.1475-4983.2006.00613.x · Cited by 267
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The host selects mucosal and luminal associations of coevolved gut microorganisms: a novel concept
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Evolutionary consequences of antibiotic use for the resistome, mobilome and microbial pangenome
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“This may contain 4–6 × 1030 cells and hold a significant proportion of the carbon, nitrogen, and phosphorus found in living things (Whitman et al., 1998).”
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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