Extremophile - Knowledge

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suggested that the landing of spacecraft may also result in interplanetary spore transfer, given that spores can survive high-velocity impact while ejected from the spacecraft onto the planet surface. This is the first study which reported that bacteria can survive in such high-velocity impact. However, the lethal impact speed is unknown, and further experiments should be done by introducing higher-velocity impact to bacterial endospores.

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of contaminated sites as some species are capable of biodegradation under conditions too extreme for classic bioremediation candidate species. Anthropogenic activity causes the release of pollutants that may potentially settle in extreme environments as is the case with tailings and sediment released

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Dai, Shang; Chen, Qi; Jiang, Meng; Wang, Binqiang; Xie, Zhenming; Yu, Ning; Zhou, Yulong; Li, Shan; Wang, Liangyan; Hua, Yuejin; Tian, Bing (September 2021). "Colonized extremophile Deinococcus radiodurans alleviates toxicity of cadmium and lead by suppressing heavy metal accumulation and improving

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is a major environmental concern associated with many metal mines. This is due to the fact that this highly acidic water can mix with groundwater, streams, and lakes. The drainage turns the pH in these water sources from a more neutral pH to a pH lower than 4. This is close to the acidity levels of

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in the evolutionary history of the planet. Some spores and cocooned bacteria samples have been dormant for more than 40 million years; extremophiles have continued to thrive in the most extreme conditions, making them one of the most abundant lifeforms. The study of extremophiles has expanded human

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were able to survive high speed impacts up to 299±28 m/s, extreme shock, and extreme deceleration. They pointed out that this feature might allow endospores to survive and to be transferred between planets by traveling within meteorites or by experiencing atmosphere disruption. Moreover, they

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Duarte, Alysson Wagner Fernandes; dos Santos, Juliana Aparecida; Vianna, Marina Vitti; Vieira, Juliana Maíra Freitas; Mallagutti, Vitor Hugo; Inforsato, Fabio José; Wentzel, Lia Costa Pinto; Lario, Luciana Daniela; Rodrigues, Andre; Pagnocca, Fernando Carlos; Pessoa Junior, Adalberto (11 December

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has been shown to effectively manage the concentration of these metals within twelve hours of introduction. Some acidophilic microorganisms are effective at metal remediation in acidic environments due to proteins found in their periplasm, not present in any mesophilic organisms, allowing them to

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at temperatures exceeding 60 °C (140 °F). The expression of this gene is a crucial precursor to the bioremediation process. Fungi that have been genetically modified with cold-adapted enzymes to tolerate differing pH levels and temperatures have been shown to be effective at remediating

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Any bacteria capable of inhabiting radioactive mediums can be classified as an extremophile. Radioresistant organisms are therefore critical in the bioremediation of radionuclides. Uranium is particularly challenging to contain when released into an environment and very harmful to both human and

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Over 65 prokaryotic species are known to be naturally competent for genetic transformation, the ability to transfer DNA from one cell to another cell followed by integration of the donor DNA into the recipient cell's chromosome. Several extremophiles are able to carry out species-specific DNA

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zone. Some microorganisms have been found thriving inside rocks up to 1,900 feet (580 m) below the sea floor under 8,500 feet (2,600 m) of ocean off the coast of the northwestern United States. According to one of the researchers, "You can find microbes everywhere—they're extremely

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There are many classes of extremophiles that range all around the globe; each corresponding to the way its environmental niche differs from mesophilic conditions. These classifications are not exclusive. Many extremophiles fall under multiple categories and are classified as

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was able to survive under Martian radiation at a wavelength that was considered extremely lethal to most bacteria. This discovery is significant because it indicates that not only bacterial spores, but also growing cells can be remarkably resistant to strong UV radiation.

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had a half life of 15 seconds under the same conditions. The catalase will have applications for removal of hydrogen peroxide in industrial processes such as pulp and paper bleaching, textile bleaching, food pasteurization, and surface decontamination of food packaging.

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Schulze-Makuch, Dirk; Haque, Shirin; De Sousa Antonio, Marina Resendes; Ali, Denzil; Hosein, Riad; Song, Young C.; Yang, Jinshu; Zaikova, Elena; Beckles, Denise M.; Guinan, Edward; Lehto, Harry J.; Hallam, Steven J. (2011). "Microbial Life in a Liquid Asphalt Desert".

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microbes metabolize hydrocarbons which assists in the remediation of hazardous, oil-based pollutants in the Arctic and Antarctic regions. These specific microbes are used in this region due to their ability to perform their functions at extremely cold temperatures.

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mediates chromosomal marker exchange with high frequency. Recombination rates exceed those of uninduced cultures by up to three orders of magnitude. Frols et al. and Ajon et al. hypothesized that cellular aggregation enhances species-specific DNA transfer between

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that might be different from that on Earth. Astrobiologists are particularly interested in studying extremophiles, as it allows them to map what is known about the limits of life on Earth to potential extraterrestrial environments For example, analogous deserts of

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ecosystem health. The NANOBINDERS project is equipping bacteria that can survive in uranium rich environments with gene sequences that enable proteins to bind to uranium in mining effluent, making it more convenient to collect and dispose of. Some examples are

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Lollar, Garnet S.; Warr, Oliver; Telling, Jon; Osburn, Magdalena R.; Lollar, Barbara Sherwood (2019). "'Follow the Water': Hydrogeochemical Constraints on Microbial Investigations 2.4 km Below Surface at the Kidd Creek Deep Fluid and Deep Life Observatory".

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Tom Gheysens from Ghent University in Belgium and some of his colleagues have presented research findings that show spores from a species of Bacillus bacteria survived and were still viable after being heated to temperatures of 420 °C (788 °F).

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cells in order to repair damaged DNA by means of homologous recombination. Van Wolferen et al. noted that this DNA exchange process may be crucial under DNA damaging conditions such as high temperatures. It has also been suggested that DNA transfer in

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was one of the bacteria which displayed not only survival but also robust cellular growth under these conditions of hyperacceleration which are usually found only in cosmic environments, such as on very massive stars or in the shock waves of

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researchers. The catalase operates over a temperature range from 30 °C to over 94 °C and a pH range from 6–10. This catalase is extremely stable compared to other catalases at high temperatures and pH. In a comparative study, the

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from deep-sea mining activity. While most bacteria would be crushed by the pressure in these environments, piezophiles can tolerate these depths and can metabolize pollutants of concern if they possess bioremediation potential.

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may be an early form of sexual interaction similar to the more well-studied bacterial transformation systems that involve species-specific DNA transfer leading to homologous recombinational repair of DNA damage (and see

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are likely not limited to Antarctica after discovering the two genes previously linked to their "atmospheric chemosynthesis" in soil of two other similar cold desert sites, which provides further information on this

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Wynn-Williams DA, Newton EM, Edwards HG (2001). "The role of habitat structure for biomolecule integrity and microbial survival under extreme environmental stress in Antarctica (and Mars?): ecology and technology".

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are hyperthermophilic archaea. Exposure of these organisms to the DNA damaging agents UV irradiation, bleomycin or mitomycin C induces species-specific cellular aggregation. UV-induced cellular aggregation of

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Heuer, Verena B.; Inagaki, Fumio; Morono, Yuki; Kubo, Yusuke; Spivack, Arthur J.; Viehweger, Bernhard; Treude, Tina; Beulig, Felix; Schubotz, Florence; Tonai, Satoshi; Bowden, Stephen A. (4 December 2020).

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which live 7,900 feet (2,400 m) below the surface, and which breathe sulfur in order to survive. These organisms are also remarkable due to eating rocks such as pyrite as their regular food source.

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battery acid or stomach acid. Exposure to the polluted water can greatly affect the health of plants, humans, and animals. However, a productive method of remediation is to introduce the extremophile,

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tolerant) archaeon, is capable of natural genetic transformation. Cytoplasmic bridges are formed between cells that appear to be used for DNA transfer from one cell to another in either direction.

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Marion, Giles M.; Fritsen, Christian H.; Eicken, Hajo; Payne, Meredith C. (December 2003). "The Search for Life on Europa: Limiting Environmental Factors, Potential Habitats, and Earth Analogues".

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and living under the Martian surface. Research indicates it is unlikely that Martian microbes exist on the surface or at shallow depths, but may be found at subsurface depths of around 100 meters.

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Good, Nathan M.; Martinez-Gomez, N. Cecilia (1 January 2021), "Expression, purification and testing of lanthanide-dependent enzymes in Methylorubrum extorquens AM1", in Cotruvo, Joseph A. (ed.),

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Glud RN, Wenzhöfer F, Middelboe M, Oguri K, Turnewitsch R, Canfield DE, Kitazato H (17 March 2013). "High rates of microbial carbon turnover in sediments in the deepest oceanic trench on Earth".

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is the multidisciplinary field that investigates the deterministic conditions and contingent events with which life arises, distributes, and evolves in the universe. Astrobiology makes use of

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gene making it mercury resistant. Industrial effluent contain high levels of metals that can be detrimental to both human and ecosystem health. In extreme heat environments the extremophile

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Barney BL, Pratt SN, Austin DE (June 2016). "Survivability of bare, individual Bacillus subtilis spores to high-velocity surface impact: Implications for microbial transfer through space".

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has great flexibility for surviving in extreme environments—niches that are acidic, extraordinarily hot, or with irregular air pressure for example—that would be completely inhospitable to

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There are multiple potential destinations for hydrocarbons after an oil spill has settled and currents routinely deposit them in extreme environments. Methane bubbles resulting from the

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mol per kilogram. The combination of low temperatures and high pressures in this environment result in low microbial activity. However, bacteria that are present including species of

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Gaudin M, Gauliard E, Schouten S, Houel-Renault L, Lenormand P, Marguet E, Forterre P (February 2013). "Hyperthermophilic archaea produce membrane vesicles that can transfer DNA".

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start to help neutralize the acidity of the waste water. This is a way to reduce the environmental impact and help remediate the damage caused by the acid mine drainage leaks.

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Ahmed I, Yokota A, Fujiwara T (March 2007). "A novel highly boron tolerant bacterium, Bacillus boroniphilus sp. nov., isolated from soil, that requires boron for its growth".

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Hong, Soon Ho; Kim, Jin Sik; Lee, Sang Yup; In, Yong Ho; Choi, Sun Shim; Rih, Jeong-Keun; Kim, Chang Hoon; Jeong, Haeyoung; Hur, Cheol Goo; Kim, Jae Jong (19 September 2004).

3327:"Soil Microbiomes With the Genetic Capacity for Atmospheric Chemosynthesis Are Widespread Across the Poles and Are Associated With Moisture, Carbon, and Nitrogen Limitation" 5136:

Gaudin M, Krupovic M, Marguet E, Gauliard E, Cvirkaite-Krupovic V, Le Cam E, Oberto J, Forterre P (April 2014). "Extracellular membrane vesicles harbouring viral genomes".

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New sub-types of extremophiles are identified frequently and the sub-category list for extremophiles is always growing. For example, microbial life lives in the liquid

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Radioresistance has also been observed in certain species of macroscopic lifeforms. The lethal dose required to kill up to 50% of a tortoise population is 40,000

1299:. This extremophile is useful for its bioleaching property. It helps to break down minerals in the waste water created by the mine. By breaking down the minerals 2260: 1495:

is one of the most radioresistant organisms known. This bacterium can also survive cold, dehydration, vacuum and acid and is thus known as a polyextremophile.

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Since the definition of an extreme environment is relative to an arbitrarily defined standard, often an anthropocentric one, these organisms can be considered

2399:"Thermococcus barophilus sp. nov., a new barophilic and hyperthermophilic archaeon isolated under high hydrostatic pressure from a deep-sea hydrothermal vent" 3805:

Chatterjee, S.K.; Bhattacharjee, I.; Chandra, G. (March 2010). "Biosorption of heavy metals from industrial waste water by Geobacillus thermodenitrificans".

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Extracellular membrane vesicles (MVs) might be involved in DNA transfer between different hyperthermophilic archaeal species. It has been shown that both

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Reddy, C. M.; Arey, J. S.; Seewald, J. S.; Sylva, S. P.; Lemkau, K. L.; Nelson, R. K.; Carmichael, C. A.; McIntyre, C. P.; Fenwick, J.; Ventura, G. T.;

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ability under particular conditions. Studying extreme environments on Earth can help researchers understand the limits of habitability on other worlds.

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are resistant to the harsh conditions of the environment and are therefore candidate species for limiting the extent of contamination of these metals.

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Valdés, Jorge; Pedroso, Inti; Quatrini, Raquel; Dodson, Robert J; Tettelin, Herve; Blake, Robert; Eisen, Jonathan A; Holmes, David S (December 2008).

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van Wolferen M, Ajon M, Driessen AJ, Albers SV (July 2013). "How hyperthermophiles adapt to change their lives: DNA exchange in extreme conditions".

2827: 4073: 2935: 1953: 1396:. Research indicates that extremophiles inhabit the asphalt lake in populations ranging between 10 and 10 cells/gram. Likewise, until recently, 5453: 1406:, borophiles came into discussion. Studying these borophiles may help illuminate the mechanisms of both boron toxicity and boron deficiency. 3712:"Volatilization of Mercury under Acidic Conditions from Mercury-polluted Soil by a Mercury-resistant Acidithiobacillus ferrooxidans SUG 2-2" 3951:

Phi, Manh Tri; Singer, Helena; Zäh, Felix; Haisch, Christoph; Schneider, Sabine; Op den Camp, Huub J. M.; Daumann, Lena J. (1 March 2024).

1618: 775:(faux Ancient Latin/Greek for 'affection for many extremes') is an organism that qualifies as an extremophile under more than one category. 4992:

Fröls S, Ajon M, Wagner M, Teichmann D, Zolghadr B, Folea M, Boekema EJ, Driessen AJ, Schleper C, Albers SV, et al. (November 2008).

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Mastascusa V, Romano I, Di Donato P, Poli A, Della Corte V, Rotundi A, Bussoletti E, Quarto M, Pugliese M, Nicolaus B (September 2014).

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Ajon M, Fröls S, van Wolferen M, Stoecker K, Teichmann D, Driessen AJ, Grogan DW, Albers SV, Schleper C, et al. (November 2011).

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Fakayode, S.O. (2005). "Impact assessment of industrial effluent on water quality of the receiving Alaro River in Ibadan, Nigeria".

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Merino, Nancy; Aronson, Heidi S.; Bojanova, Diana P.; Feyhl-Buska, Jayme; Wong, Michael L.; Zhang, Shu; Giovannelli, Donato (2019).

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Merino, Nancy; Aronson, Heidi S.; Bojanova, Diana P.; Feyhl-Buska, Jayme; Wong, Michael L.; Zhang, Shu; Giovannelli, Donato (2019).

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Neufeld, Josh; Clarke, Andrew; Morris, G. John; Fonseca, Fernanda; Murray, Benjamin J.; Acton, Elizabeth; Price, Hannah C. (2013).

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Rosenshine I, Tchelet R, Mevarech M (September 1989). "The mechanism of DNA transfer in the mating system of an archaebacterium".

5986: 5488: 4264:"Bioremediation of oil spills: An annotated selection of World Wide Web sites relevant to the topics in Microbial Biotechnology" 4199:"Flavobacterium petrolei sp. nov., a novel psychrophilic, diesel-degrading bacterium isolated from oil-contaminated Arctic soil" 4706: 4542: 5380: 4781: 4338: 4017: 2802: 2234: 194: 3597:

Margesin, R.; Schinner, F. (1 September 2001). "Biodegradation and bioremediation of hydrocarbons in extreme environments".

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and further strengthens the extremophile evidence that supports the potential existence of microbial life on alien planets.

4994:"UV-inducible cellular aggregation of the hyperthermophilic archaeon Sulfolobus solfataricus is mediated by pili formation" 1477:

enzymes used in clinical diagnostics and starch liquefaction are produced commercially by several biotechnology companies.

101:, i.e., environments with conditions approaching or stretching the limits of what known life can adapt to, such as extreme 2963:"The adaptation potential of extremophiles to Martian surface conditions and its implication for the habitability of Mars" 603:, which has flat square-shaped cells with gas vesicles that allow flotation to the surface, most likely to acquire oxygen. 5961: 5224:"Insights into dynamics of mobile genetic elements in hyperthermophilic environments from five new Thermococcus plasmids" 3767:

Nagajyoti, P.C. (2008). "Heavy metal toxicity: Industrial Effluent Effect on Groundnut (Arachis hypogaea L.) Seedlings".

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Marteinsson VT, Birrien JL, Reysenbach AL, Vernet M, Marie D, Gambacorta A, Messner P, Sleytr UB, Prieur D (April 1999).

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Takeuchi, Fumiaki; Iwahori, Kenji; Kamimura, Kazuo; Negishi, Atsunori; Maeda, Terunobu; Sugio, Tsuyoshi (January 2001).

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were found to be capable of bioremediation, albeit at a tenth of the speed they would perform at sea level pressure.

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found microorganisms in 120 °C (248 °F) sediment that is 1.2 km (0.75 mi) below seafloor in the

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can be transferred via MVs. Notably, a horizontal plasmid transfer has been documented between hyperthermophilic

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transfer, as described below. However, it is not yet clear how common such a capability is among extremophiles.

1177: 3850:"Periplasmic proteins of the extremophile Acidithiobacillus ferrooxidans: a high throughput proteomics analysis" 3403: 4048: 1427: 1255: 1242: 32: 5364:

Encyclopedia of Earth, National Council of Science & the Environment, Eds. E. Monosson & C. Cleveland

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On 20 August 2014, scientists confirmed the existence of microorganisms living half a mile below the ice of

3432:"DNA Damage and Survival Time Course of Deinococcal Cell Pellets During 3 Years of Exposure to Outer Space" 1436:, which initiates the breakdown of hydrogen peroxide into oxygen and water, was isolated from an organism, 5422: 1100: 5481: 3540:"Composition and fate of gas and oil released to the water column during the Deepwater Horizon oil spill" 2147: 1151: 1122: 1014: 658:

An organism with optimal growth at a concentration of dissolved salts of 50 g/L (= 5% m/v) or above.

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An organism with optimal growth conditions in high concentrations of carbon dioxide. An example would be

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Exo-/Astro-biology: Proceedings of the First European Workshop, 21–23 May 2001, ESRIN, Fracscati, Italy

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tolerance was unknown, but a strong borophile was discovered in bacteria. With the recent isolation of

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being subject to conditions of extreme gravity. The bacteria were cultivated while being rotated in an

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Ray, Angelique E.; Zhang, Eden; Terauds, Aleks; Ji, Mukan; Kong, Weidong; Ferrari, Belinda C. (2020).

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2017). "Cold-adapted enzymes produced by fungi from terrestrial and marine Antarctic environments".

3404:"Bacteria from Earth can survive in space and could endure the trip to Mars, according to new study" 3274: 5981: 5643: 4840:"Transformation in Micrococcus radiodurans and the ultraviolet sensitivity of its transforming DNA" 2436:"Diversity and phylogenetic profiling of niche-specific Bacilli from extreme environments of India" 1365:, compared to only 800 roentgens needed to kill 50% of a human population. In experiments exposing 1248: 938: 571: 205:

adaptable to conditions, and survive wherever they are." A key to extremophile adaptation is their

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that have been found that are 40 million years old on Earth—and we know they're very hardened to

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catalase exhibited a half life of 15 days at 80 °C and pH 10 while a catalase derived from

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Chandna, S.; Dwarakanath, B. S.; Seth, R. K.; Khaitan, D.; Adhikari, J. S.; Jain, V. (2004).

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and higher doses, in contrast with human cells that showed similar damage at only 2 Gy.

1344: 1038: 599: 145: 130: 4140:"Acidithiobacillus ferrooxidans metabolism: from genome sequence to industrial applications" 3252: 1952: 5674: 5235: 5188: 5145: 4955: 4851: 4673: 4578: 4362: 4210: 3909: 3248: 3195: 3118: 3015: 2977: 2862: 2790: 2703: 2642: 2575: 2351: 2304: 2061: 2018: 1741: 1509: 1096: 756: 669: 282: 98: 5359: 2532: 1919: 1260: 185:

were found living in the cold and dark in a lake buried a half-mile deep under the ice in

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Deguchi S, Shimoshige H, Tsudome M, Mukai SA, Corkery RW, Ito S, Horikoshi K (May 2011).

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were found 1.1 kilometers below water surface level and at concentrations as high as 183

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Heising-Goodman, Carolyn (March 1981). "Nuclear Power and Its Environmental Effects".

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Kim W, Tengra FK, Young Z, Shong J, Marchand N, Chan HK, et al. (29 April 2013).

2477:"The genome sequence of the capnophilic rumen bacterium Mannheimia succiniciproducens" 2049: 1800: 1025:

in ways "not observed on Earth" and in ways that "can lead to increases in growth and

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Wilson ZE, Brimble MA (January 2009). "Molecules derived from the extremes of life".

5263: 5204: 5200: 5161: 5107: 5064: 5059: 5015: 5010: 4971: 4928: 4879: 4820: 4777: 4693: 4637: 4594: 4517: 4437: 4419: 4398:"Extremophilic Microfactories: Applications in Metal and Radionuclide Bioremediation" 4378: 4334: 4283: 4279: 4244: 4226: 4179: 4161: 4067: 4023: 4013: 3980: 3972: 3933: 3925: 3881: 3830: 3822: 3741: 3733: 3684: 3676: 3622: 3614: 3579: 3561: 3535: 3514: 3477: 3465: 3378: 3366: 3348: 3221: 3136: 3043: 2890: 2798: 2760: 2737: 2719: 2668: 2593: 2506: 2498: 2416: 2379: 2320: 2261:"Turn Up the Heat: Bacterial Spores Can Take Temperatures in the Hundreds of Degrees" 2205: 2107: 2095: 2087: 2009: 1865: 1834:"Living at the Extremes: Extremophiles and the Limits of Life in a Planetary Context" 1814: 1757: 1708: 1696: 1665:"Living at the Extremes: Extremophiles and the Limits of Life in a Planetary Context" 1457: 1180:

increase in solubility and bioavailability with increasing temperature. Thermophilic

1087: 1051: 1022: 979: 887: 879: 801: 585: 493: 466: 371: 258: 245: 167: 5076: 4606: 4529: 4113: 3634: 3501:

Frid, Christopher L. J.; Caswell, Bryony A. (23 November 2017). "Marine Pollution".

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An organism with optimal growth at temperatures of 15 °C (59 °F) or lower.

5976: 5928: 5812: 5610: 5605: 5581: 5336: 5328: 5295: 5253: 5243: 5196: 5153: 5119: 5099: 5054: 5005: 4963: 4918: 4910: 4869: 4859: 4810: 4681: 4649: 4629: 4586: 4509: 4498:"Radiation responses of Sf9, a highly radioresistant lepidopteran insect cell line" 4427: 4409: 4370: 4329: 4324: 4316: 4291: 4275: 4234: 4218: 4169: 4151: 4005: 3964: 3953:"Assessing Lanthanide-Dependent Methanol Dehydrogenase Activity: The Assay Matters" 3917: 3871: 3861: 3814: 3753: 3723: 3696: 3666: 3658: 3606: 3569: 3551: 3506: 3455: 3445: 3356: 3338: 3256: 3211: 3203: 3184:"Extremophiles survival to simulated space conditions: an astrobiology model study" 3126: 3072: 3067: 3033: 3023: 2880: 2870: 2727: 2711: 2658: 2650: 2583: 2488: 2461: 2447: 2406: 2369: 2359: 2312: 2195: 2185: 2077: 2069: 2026: 1855: 1845: 1804: 1796: 1749: 1686: 1676: 1227:

are highly oxidative environments that can produce high levels of lead or cadmium.

960: 947: 932: 677: 559:. Polyextremophiles are well known for their ability to tolerate both high and low 460: 326: 4914: 4685: 4088: 3818: 3662: 2050:"Temperature limits to deep subseafloor life in the Nankai Trough subduction zone" 1769: 1081:

In June 2016, scientists from Brigham Young University conclusively reported that

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Rossi M, Ciaramella M, Cannio R, Pisani FM, Moracci M, Bartolucci S (July 2003).

5248: 5040:"UV-inducible DNA exchange in hyperthermophilic archaea mediated by type IV pili" 4815: 4798: 4713: 4513: 4197:

Chaudhary, Dhiraj Kumar; Kim, Dong-Uk; Kim, Dockyu; Kim, Jaisoo (11 March 2019).

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Capable of tolerating high levels of dissolved heavy metals in solution, such as

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has been shown to be effective in remediating mercury in acidic soil due to its

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An organism with optimal growth in environments with a high sugar concentration.

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Proceedings of the National Academy of Sciences of the United States of America

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Rare-Earth Element Biochemistry: Methanol Dehydrogenases and Lanthanide Biology

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Proceedings of the National Academy of Sciences of the United States of America

2692:"Niche partitioning of diverse sulfur-oxidizing bacteria at hydrothermal vents" 2434:

Yadav AN, Verma P, Kumar M, Pal KK, Dey R, Gupta A, et al. (31 May 2014).

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species have demonstrated higher gene expression for the alkane mono-oxygenase

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An organism with optimal growth at temperatures above 45 °C (113 °F).

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An organism with optimal growth at temperatures above 80 °C (176 °F).

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Strange life-forms found deep in a mine point to vast 'underground Galapagos'

4423: 4414: 4382: 4287: 4230: 4165: 3976: 3929: 3826: 3737: 3680: 3618: 3565: 3450: 3352: 3343: 2723: 2687: 2559: 2502: 2121: 2091: 1850: 1681: 871: 308: 249: 198: 5414: 5157: 4967: 4497: 4156: 3952: 3556: 2875: 2654: 2588: 2563: 2073: 1046:. It is not currently known if such resistant microbes could have withstood 594: 137:

of environments made hazardous to humans due to pollution or contamination.

5901: 5862: 5837: 5817: 5656: 5591: 5350: 5307: 5267: 5208: 5165: 5111: 5068: 5019: 4824: 4641: 4598: 4521: 4441: 4248: 4183: 4027: 3984: 3968: 3937: 3885: 3834: 3745: 3688: 3626: 3583: 3469: 3370: 3225: 3140: 3047: 2894: 2741: 2672: 2597: 2510: 2420: 2383: 2324: 2209: 2099: 1869: 1818: 1761: 1700: 1635: 1613: 1576: 1366: 1311: 1111:

The same month, scientists reported that bacteria from Earth, particularly

964: 908: 855: 780: 576: 552: 403: 320: 4975: 4932: 4883: 4864: 4590: 3610: 2715: 2190: 898:

to investigate the possibility of life on other worlds and help recognize

5868: 5822: 5601: 5586: 5516: 4374: 4004:, Methods in Enzymology, vol. 650, Academic Press, pp. 97–118, 2151: 1986: 1629: 1504: 1377: 1224: 1160: 1118: 1105: 1010: 827: 710: 622: 580: 474: 389: 377: 350: 102: 45: 3728: 3711: 3671: 2910:"Harvard study suggests asteroids might play key role in spreading life" 2621:"Absence of detectable arsenate in DNA from arsenate-grown GFAJ-1 cells" 2619:

Reaves ML, Sinha S, Rabinowitz JD, Kruglyak L, Redfield RJ (July 2012).

759:

above 10 MPa (= 99 atm = 1,450 psi). Also referred to as

5883: 5712: 5576: 5566: 5556: 5551: 5521: 5511: 1640: 1393: 1126: 1061: 987: 968: 904: 750: 730: 663: 636: 556: 435: 429: 363: 290: 206: 201: 186: 5432:

DaveDarling's Encyclopedia of Astrobiology, Astronomy, and Spaceflight

4557: 2690:; Vidoudez, Charles; Amann, Rudolf; Meyerdierks, Anke (4 April 2017). 2082: 735:

An organism with optimal growth in nutritionally limited environments.

166:. Some scientists even concluded that life may have begun on Earth in 5736: 5615: 5571: 5531: 5299: 4799:"Natural genetic transformation: prevalence, mechanisms and function" 4707:

World’s Oldest Groundwater Supports Life Through Water-Rock Chemistry

3300:"Bacteria that "eat" only air found in cold deserts around the world" 2030: 1753: 1623: 1410: 1362: 1166: 1082: 1043: 1026: 956: 899: 891: 867: 863: 837: 812:

An organism with optimal growth conditions in high concentrations of

760: 740: 653: 548: 296: 178: 174: 110: 82: 3131: 3106: 2686:

Meier, Dimitri V; Pjevac, Petra; Bach, Wolfgang; Hourdez, Stephane;

2493: 2476: 1729: 1306: 822:, a sulfur-oxidizing bacteria that inhabits deep-water sulfur vents. 133:. Extremophiles are also of interest because of their potential for 5935: 5661: 5634: 5536: 5526: 5178: 3408: 1526: 1473: 1433: 1199:

hydrocarbon contamination in freezing conditions in the Antarctic.

1188: 1033: 1018: 927: 916: 182: 118: 114: 106: 94: 90: 4573: 3004:"Spaceflight promotes biofilm formation by Pseudomonas aeruginosa" 2637: 2396: 5697: 1879:

Material was copied from this source, which is available under a

1567: 1389: 1369: 1182: 895: 883: 875: 859: 701: 697: 173:

According to astrophysicist Steinn Sigurdsson, "There are viable

5466: 1831: 1662: 5687: 3160: 2564:"GFAJ-1 is an arsenate-resistant, phosphate-dependent organism" 1571: 1279:

This is a before and after picture of a cleanup project of the

1172: 975: 813: 722: 693: 648:

a bacterium that inhabits a ruminant animal's digestive system.

540:. A polyextremophile living at the summit of a mountain in the 314: 129:

knowledge of the limits of life, and informs speculation about

5419: 5089: 3804: 3709: 5436: 5222:

Krupovic M, Gonnet M, Hania WB, Forterre P, Erauso G (2013).

5135: 5037: 4896: 4495: 3483: 3384: 3181: 2848: 2618: 1875: 1397: 923: 412: 367: 3647: 5314: 5221: 4137: 2006: 1194: 1006: 912: 705: 334: 4897:

Koyama Y, Hoshino T, Tomizuka N, Furukawa K (April 1986).

4796: 3788:

African Journal of Environmental Assessment and Management

2851:"Microbial growth at hyperaccelerations up to 403,627 x g" 2779: 2557: 2337: 994:

in the Mars Simulation Laboratory (MSL) maintained by the

193:, the deepest place in Earth's oceans. Expeditions of the 4945: 4463:. Vol. 171, no. 21. p. 325. Archived from 1943:"Bacteria Found Deep Under Antarctic Ice, Scientists Say" 5456:

Encyclopedia of Astrobiology, Astronomy, and Spaceflight

5398:

Extreme Environments - Science Education Resource Center

4991: 4457:"Dark Power: Pigment seems to put radiation to good use" 4049:"Actions Eliminate Long-Time, Major Acid Mine Discharge" 2294: 1068: 5370: 4797:

Johnsborg O, Eldholm V, Håvarstein LS (December 2007).

4662: 2172:

Reed CJ, Lewis H, Trejo E, Winston V, Evilia C (2013).

950:(i.e. 403,627 times the gravity experienced on Earth). 628: 614: 560: 301: 5373:

Polyextremophiles: life under multiple forms of stress

3489:

Creative Commons Attribution 4.0 International License

3390:

Creative Commons Attribution 4.0 International License

2685: 2046: 1920:"Impacts 'more likely' to have spread life from Earth" 1881:

Creative Commons Attribution 4.0 International License

1720: 1718: 1626:, an organism that grows best in moderate temperatures 1376:, significant DNA damage was detected only at 20 1095:

In August 2020 scientists reported that bacteria that

967:. The research has implications on the feasibility of 254:

110 °C (230 °F) to 121 °C (250 °F)

5912: 3533: 2171: 278:−20 °C (−4 °F) to −25 °C (−13 °F) 4479:"Tortoise blood fights radiation sickness - UPI.com" 4448: 3156:"Life Confirmed Under Antarctic Ice; Is Space Next?" 1783:

Shukla, Awadhesh Kumar; Singh, Amit Kishore (2020).

1724: 1517:

species are also capable of genetic transformation.

1223:

protect themselves from high proton concentrations.

672:

above 50 MPa (= 493 atm = 7,252 psi).

4619: 3001: 2820:"Bacteria Grow Under 400,000 Times Earth's Gravity" 1715: 1632:, an organism that grows best in a neutral pH level 1032:On 19 May 2014, scientists announced that numerous 27:

Organisms capable of living in extreme environments

4196: 3997: 3950: 2433: 2331: 2258: 2216: 3324: 3238: 3107:"Lakes under the ice: Antarctica's secret garden" 2927: 1307:Oil-based, hazardous pollutants in Arctic regions 5948: 5371:Seckbach J, Oren A, Stan-Lotter H, eds. (2013). 4728:, By Corey S. Powell, 7 Sept. 2019, nbcnews.com. 3898: 3596: 2403:International Journal of Systematic Bacteriology 2148:"Intraterrestrials: Life Thrives in Ocean Floor" 1137:Extremophiles can also be useful players in the 922:Recent research carried out on extremophiles in 579:(salinity> 20% w/v), in which the eukaryotic 5357: 5131: 5129: 4352: 3544:Proceedings of the National Academy of Sciences 3430:Kawaguchi, Yuko; et al. (26 August 2020). 3429: 2174:"Protein adaptations in archaeal extremophiles" 1383: 3188:Origins of Life and Evolution of the Biosphere 1586:species, respectively belonging to the orders 1042:, may be resistant to methods usually used in 978:survived and showed remarkable results on the 158:In the 1980s and 1990s, biologists found that 5482: 5285: 4837: 4770:Extremophiles: Microbiology and Biotechnology 3060: 2995: 2960: 2826:. National Geographic Society. Archived from 2290: 2288: 2286: 2284: 2282: 2280: 2278: 2276: 2274: 2259:Magazine, Smithsonian; Schulze-Makuch, Dirk. 1976: 1974: 1972: 1894:"Mars Exploration Rover Launches – Press kit" 5126: 4987: 4985: 4454: 3395: 3275:"Microbes living on air a global phenomenon" 3147: 3098: 3054: 2954: 2754: 2474: 1619:List of microorganisms tested in outer space 1240:on their biological processes. For example, 5437:The International Society for Extremophiles 5033: 5031: 5029: 4745:. U.S. Department of Energy. Archived from 4072:: CS1 maint: numeric names: authors list ( 3716:Bioscience, Biotechnology, and Biochemistry 3500: 3423: 2558:Erb TJ, Kiefer P, Hattendorf B, Günther D, 2340:"A Low Temperature Limit for Life on Earth" 2139: 1934: 1782: 1609:Dissimilatory metal-reducing microorganisms 1117:, were found to survive for three years in 974:On 26 April 2012, scientists reported that 5489: 5475: 4502:International Journal of Radiation Biology 3401: 2271: 2122:"T-Limit of the Deep Biosphere off Muroto" 2000: 1983:"Microbes Thrive in Deepest Spot on Earth" 1969: 593:is grown commercially for the carotenoid, 151:Diversity of extreme environments on Earth 5340: 5257: 5247: 5058: 5009: 4982: 4922: 4873: 4863: 4814: 4761: 4572: 4543:Microbial Life Found in Hydrocarbon Lake. 4431: 4413: 4328: 4295: 4238: 4173: 4155: 3875: 3865: 3766: 3727: 3670: 3573: 3555: 3459: 3449: 3360: 3342: 3215: 3130: 3037: 3027: 2970:EGU General Assembly Conference Abstracts 2884: 2874: 2817: 2731: 2662: 2636: 2587: 2492: 2451: 2410: 2373: 2363: 2199: 2189: 2165: 2081: 1917: 1859: 1849: 1825: 1808: 1690: 1680: 963:is essential for successful growth under 959:. Analysis showed that the small size of 5026: 4838:Moseley BE, Setlow JK (September 1968). 3785: 3063:"Microbial stowaways to Mars identified" 2936:"Lichen survives harsh Mars environment" 1318: 1274: 946:at high speeds corresponding to 403,627 570: 144: 31: 4767: 4739:"Bioenergy and Industrial Microbiology" 4395: 4261: 2933: 1125:. These findings support the notion of 575:Microscopic image from the hypersaline 93:that is able to live (or in some cases 14: 5949: 4086: 4046: 3599:Applied Microbiology and Biotechnology 3487:Text and images are available under a 3388:Text and images are available under a 2961:De Vera JP, Kohler U (26 April 2012). 2907: 2525: 1940: 1658: 1656: 796:Organisms resistant to high levels of 5470: 4311:"Toxicological Profile for Uranium", 3402:Strickland, Ashley (26 August 2020). 2145: 2042: 2040: 1728:; Mancinelli, Rocco (February 2001). 1270: 195:International Ocean Discovery Program 5454:Polyextremophile on David Darling's 5420:The Research Center of Extremophiles 4455:Castelvecchi, Davide (26 May 2007). 4396:Marques, Catarina R. (1 June 2018). 3769:Journal of Applied Sciences Research 3153: 1980: 1409:In July 2019, a scientific study of 1236:Some bacteria are known to also use 4047:US EPA, REG 03 (9 September 2016). 3854:Molecular & Cellular Proteomics 3847: 3104: 1653: 1067:In September 2015, scientists from 842:An organism with optimal growth at 755:An organism with optimal growth in 668:An organism with optimal growth at 627:An organism with optimal growth at 613:An organism with optimal growth at 24: 5415:Eukaryotes in extreme environments 5278: 5181:Environmental Microbiology Reports 3511:10.1093/oso/9780198726289.001.0001 2757:The life and death of planet Earth 2533:"Studies refute arsenic bug claim" 2233:. 2015. p. 59. Archived from 2037: 850: 526: 140: 25: 5998: 5808:Acidophiles in acid mine drainage 5496: 5391: 4262:Wackett, Lawrence P. (May 2012). 3651:Critical Reviews in Biotechnology 1801:10.2174/1389202921999200422122253 1325:Uranium § Biotic and abiotic 1132: 589:, can be tentatively identified. 170:far beneath the ocean's surface. 5934: 5922: 5333:10.1128/JB.185.13.3683-3689.2003 5201:10.1111/j.1758-2229.2012.00348.x 5060:10.1111/j.1365-2958.2011.07861.x 5011:10.1111/j.1365-2958.2008.06459.x 4280:10.1111/j.1751-7915.2011.00330.x 4087:US EPA, OW (15 September 2015). 3482: 3383: 2824:National Geographic – Daily News 1874: 1421: 1178:Polycyclic aromatic hydrocarbons 1003:Rensselaer Polytechnic Institute 1001:On 29 April 2013, scientists at 819:Sulfurovum epsilonproteobacteria 5215: 5172: 5083: 4939: 4890: 4831: 4790: 4731: 4719: 4716:, 29 July 2019, deepcarbon.net. 4700: 4656: 4613: 4551: 4536: 4489: 4471: 4389: 4346: 4304: 4255: 4190: 4131: 4106: 4080: 4040: 3991: 3944: 3892: 3841: 3798: 3779: 3760: 3703: 3641: 3590: 3527: 3494: 3318: 3292: 3267: 3232: 3175: 3061:Madhusoodanan J (19 May 2014). 2938:. Skymania News. Archived from 2901: 2842: 2811: 2773: 2748: 2679: 2612: 2551: 2468: 2427: 2390: 2252: 2114: 1480: 1263:as cofactors to increase their 1219:Geobacillus thermodenitrificans 1145: 1044:spacecraft assembly clean rooms 5987:Microbial growth and nutrition 4315:, CRC Press, 12 January 2002, 4313:ATSDR's Toxicological Profiles 3807:Journal of Hazardous Materials 1911: 1886: 1776: 1730:"Life in extreme environments" 1465:DNA modifying enzymes such as 1428:Extremophiles in biotechnology 1256:Methylobacterium radiotolerans 1243:Methylacidiphilum fumariolicum 223:Limits of known life on Earth 13: 1: 4915:10.1128/jb.166.1.338-340.1986 4686:10.1080/01490451.2019.1641770 3900:antioxidant system in rice". 3819:10.1016/j.jhazmat.2009.09.136 3663:10.1080/07388551.2017.1379468 2908:Reuell, Peter (8 July 2019). 1646: 1356:Chernobyl Nuclear Power Plant 1209:Acidithiubacillus ferroxidans 1069:CNR-National Research Council 644:Mannheimia succiniciproducens 209:composition, affecting their 5249:10.1371/journal.pone.0049044 4816:10.1016/j.resmic.2007.09.004 4514:10.1080/09553000410001679794 3922:10.1016/j.envpol.2021.117127 3029:10.1371/journal.pone.0062437 2755:Ward PD, Brownlee D (2004). 2365:10.1371/journal.pone.0066207 2224:"NASA Astrobiology Strategy" 1941:Gorman J (6 February 2013). 1918:BBC Staff (23 August 2011). 1384:Examples and recent findings 1259:are known to be able to use 246:Submarine hydrothermal vents 68: 'extreme' and 7: 4321:10.1201/9781420061888_ch157 3241:Planetary and Space Science 2934:Baldwin E (26 April 2012). 2818:Than, Ker (25 April 2011). 1597: 1152:Deepwater Horizon oil spill 1123:International Space Station 1015:International Space Station 504:Accessed in some gold mines 10: 6003: 5962:Environmental microbiology 5793:Abiogenic petroleum origin 5726:Thermococcus gammatolerans 5138:Environmental Microbiology 4223:10.1038/s41598-019-40667-7 4010:10.1016/bs.mie.2021.02.001 3867:10.1074/mcp.M700042-MCP200 2317:10.1089/153110703322736105 1425: 1415:sulfur-breathing organisms 1322: 1121:, based on studies on the 520:, unidentified arthropods 396:Thermococcus gammatolerans 75: 5785: 5735: 5696: 5633: 5624: 5504: 5442:Idaho National Laboratory 5104:10.1007/s00792-013-0552-6 4743:Idaho National Laboratory 4712:10 September 2019 at the 4634:10.1007/s00792-006-0027-0 4402:Frontiers in Microbiology 4089:"Abandoned Mine Drainage" 3503:Oxford Scholarship Online 3437:Frontiers in Microbiology 3331:Frontiers in Microbiology 3261:10.1016/j.pss.2016.02.010 3208:10.1007/s11084-014-9397-y 3154:Mack E (20 August 2014). 3077:10.1038/nature.2014.15249 2453:10.1007/s13213-014-0897-9 2412:10.1099/00207713-49-2-351 2146:Oskin B (14 March 2013). 1981:Choi CQ (17 March 2013). 1838:Frontiers in Microbiology 1669:Frontiers in Microbiology 1604:Earliest known life forms 1561:Transformation (genetics) 1540:Sulfolobus acidocaldarius 1525:, an extreme halophilic ( 1448:Idaho National Laboratory 1444:Yellowstone National Park 1339:Geobacter metallireducens 1301:Thiobacillus ferrooxidans 1202: 718:Cupriavidus metallidurans 381: 222: 48:, a type of extremophile. 42:Yellowstone National Park 5644:Chloroflexus aurantiacus 5358:C.Michael Hogan (2010). 4803:Research in Microbiology 4768:Anitori RP, ed. (2012). 4415:10.3389/fmicb.2018.01191 3451:10.3389/fmicb.2020.02050 3344:10.3389/fmicb.2020.01936 1851:10.3389/fmicb.2019.00780 1682:10.3389/fmicb.2019.00780 1499:is competent to perform 1471:DNA polymerase and some 1249:Methylorubrum extorquens 1057:now on the planet Mars. 1009:, reported that, during 939:Paracoccus denitrificans 566: 5767:Halicephalobus mephisto 5760:Paralvinella sulfincola 5746:Cyanidioschyzon merolae 5651:Deinococcus radiodurans 5447:18 October 2014 at the 5425:11 January 2016 at the 5408:18 October 2014 at the 5375:. Dordrecht: Springer. 5321:Journal of Bacteriology 5288:Natural Product Reports 5158:10.1111/1462-2920.12235 4968:10.1126/science.2818746 4903:Journal of Bacteriology 4666:Geomicrobiology Journal 4330:2027/mdp.39015032949136 4268:Microbial Biotechnology 4157:10.1186/1471-2164-9-597 3902:Environmental Pollution 3557:10.1073/pnas.1101242108 3253:2016P&SS..125...20B 2876:10.1073/pnas.1018027108 2759:. New York: Owl Books. 2655:10.1126/science.1219861 2589:10.1126/science.1218455 2405:. 49 Pt 2 (2): 351–59. 2074:10.1126/science.abd7934 1534:Sulfolobus solfataricus 1492:Deinococcus radiodurans 1432:The thermoalkaliphilic 1333:Shewanella putrefaciens 1230:Deinococcus radiodurans 1114:Deinococcus radiodurans 1071:of Italy reported that 1050:and are present on the 996:German Aerospace Center 984:photosynthetic activity 907:are exposed to harmful 631:levels of 9.0 or above. 617:levels of 3.0 or below. 538:Thermococcus barophilus 511:Halicephalobus mephisto 447:High salt concentration 384:Deinococcus radiodurans 5047:Molecular Microbiology 4998:Molecular Microbiology 4774:Caister Academic Press 4546:the physics arXiv blog 3969:10.1002/cbic.202300811 2440:Annals of Microbiology 1522:Halobacterium volcanii 1501:genetic transformation 1283: 1281:Little Conemaugh River 1265:methanol dehydrogenase 926:involved a variety of 917:endolithic communities 816:. An example would be 604: 517:Mylonchulus brachyurus 489:~60% relative humidity 155: 62: 49: 38:Grand Prismatic Spring 5753:Galdieria sulphuraria 5682:Spirochaeta americana 5403:Extremophile Research 4865:10.1073/pnas.61.1.176 4591:10.1089/ast.2010.0488 3611:10.1007/s002530100701 3105:Fox D (August 2014). 2716:10.1038/ismej.2017.37 1413:in Canada discovered 1403:Bacillus boroniphilus 1345:Burkholderia fungorum 1319:Radioactive materials 1278: 1039:Tersicoccus phoenicis 1021:seem to adapt to the 952:P. denitrificans 757:hydrostatic pressures 670:hydrostatic pressures 600:Haloquadratum walsbyi 574: 148: 131:extraterrestrial life 126:ecologically dominant 36:The bright colors of 35: 5675:Thermus thermophilus 5317:"Extremophiles 2002" 4375:10.13182/nt81-a32724 4114:"Acid Mine Drainage" 2481:Nature Biotechnology 2265:Smithsonian Magazine 1510:Thermus thermophilus 1342:and some strains of 1099:air discovered 2017 1074:S. soflataricus 283:Rhodotorula glutinis 231:Environment / source 99:extreme environments 89: 'love') is an 5874:Radiotrophic fungus 5851:Helaeomyia petrolei 5798:Acidithiobacillales 5707:Pyrococcus furiosus 5240:2013PLoSO...849044K 5193:2013EnvMR...5..109G 5150:2014EnvMi..16.1167G 4960:1989Sci...245.1387R 4856:1968PNAS...61..176M 4678:2019GmbJ...36..859L 4583:2011AsBio..11..241S 4367:1981NucTe..52..445H 4215:2019NatSR...9.4134C 3914:2021EPoll.28417127D 3729:10.1271/bbb.65.1981 3200:2014OLEB...44..231M 3123:2014Natur.512..244F 3020:2013PLoSO...862437K 2982:2012EGUGA..14.2113D 2942:on 12 November 2020 2867:2011PNAS..108.7997D 2795:2001ESASP.496..225W 2708:2017ISMEJ..11.1545M 2647:2012Sci...337..470R 2580:2012Sci...337..467E 2356:2013PLoSO...866207C 2309:2003AsBio...3..785M 2240:on 22 December 2016 2191:10.1155/2013/373275 2066:2020Sci...370.1230H 2023:2013NatGe...6..284G 1746:2001Natur.409.1092R 1740:(6823): 1092–1101. 1497:D. radiodurans 1238:rare earth elements 980:adaptation capacity 708:. Examples include 484:McMurdo Dry Valleys 265:Pyrococcus furiosus 5462:T-Limit Expedition 4749:on 18 October 2014 4355:Nuclear Technology 4203:Scientific Reports 3536:Van Mooy, B. A. S. 1948:The New York Times 1583:Methanocaldococcus 1513:and other related 1439:Thermus brockianus 1352:Radiotrophic fungi 1287:Acid mine drainage 1284: 1271:Acid mine drainage 992:Martian conditions 798:ionizing radiation 605: 359:Ionizing radiation 342:acid mine drainage 340:Volcanic springs, 168:hydrothermal vents 156: 50: 44:, are produced by 5910: 5909: 5857:Hydrothermal vent 5781: 5780: 5719:Pyrolobus fumarii 5668:Thermus aquaticus 5382:978-94-007-6488-0 5327:(13): 3683–3689. 4954:(4924): 1387–89. 4783:978-1-904455-98-1 4467:on 24 April 2008. 4340:978-1-4200-6188-8 4019:978-0-12-823856-1 3963:(5): e202300811. 2861:(19): 7997–8002. 2804:978-92-9092-806-5 2126:www.jamstec.go.jp 2060:(6521): 1230–34. 2010:Nature Geoscience 1957:on 1 January 2022 1899:. NASA. June 2003 1546:S. acidocaldarius 1458:Aspergillus niger 1088:Bacillus subtilis 1023:space environment 990:of 34 days under 961:prokaryotic cells 888:planetary science 880:molecular biology 802:nuclear radiation 591:Dunaliella salina 586:Dunaliella salina 534:polyextremophiles 524: 523: 494:Chroococcidiopsis 467:Dunaliella salina 372:radioactive decay 259:Pyrolobus fumarii 164:complex organisms 16:(Redirected from 5994: 5939: 5938: 5927: 5926: 5925: 5918: 5813:Archaeoglobaceae 5786:Related articles 5631: 5630: 5611:Thermoacidophile 5606:Hyperthermophile 5582:Polyextremophile 5491: 5484: 5477: 5468: 5467: 5386: 5367: 5354: 5344: 5311: 5300:10.1039/b800164m 5272: 5271: 5261: 5251: 5219: 5213: 5212: 5176: 5170: 5169: 5133: 5124: 5123: 5087: 5081: 5080: 5062: 5044: 5035: 5024: 5023: 5013: 4989: 4980: 4979: 4943: 4937: 4936: 4926: 4894: 4888: 4887: 4877: 4867: 4835: 4829: 4828: 4818: 4794: 4788: 4787: 4765: 4759: 4758: 4756: 4754: 4735: 4729: 4723: 4717: 4704: 4698: 4697: 4660: 4654: 4653: 4617: 4611: 4610: 4576: 4555: 4549: 4540: 4534: 4533: 4493: 4487: 4486: 4475: 4469: 4468: 4452: 4446: 4445: 4435: 4417: 4393: 4387: 4386: 4350: 4344: 4343: 4332: 4308: 4302: 4301: 4299: 4259: 4253: 4252: 4242: 4194: 4188: 4187: 4177: 4159: 4135: 4129: 4128: 4126: 4124: 4110: 4104: 4103: 4101: 4099: 4084: 4078: 4077: 4071: 4063: 4061: 4059: 4044: 4038: 4037: 4036: 4034: 3995: 3989: 3988: 3948: 3942: 3941: 3896: 3890: 3889: 3879: 3869: 3848:Chi, A. (2007). 3845: 3839: 3838: 3802: 3796: 3795: 3783: 3777: 3776: 3764: 3758: 3757: 3731: 3707: 3701: 3700: 3674: 3645: 3639: 3638: 3594: 3588: 3587: 3577: 3559: 3550:(50): 20229–34. 3538:(18 July 2011). 3531: 3525: 3524: 3498: 3492: 3486: 3481: 3463: 3453: 3427: 3421: 3420: 3418: 3416: 3399: 3393: 3387: 3382: 3364: 3346: 3322: 3316: 3315: 3313: 3311: 3306:. 19 August 2020 3296: 3290: 3289: 3287: 3285: 3271: 3265: 3264: 3236: 3230: 3229: 3219: 3179: 3173: 3172: 3170: 3168: 3151: 3145: 3144: 3134: 3117:(7514): 244–46. 3102: 3096: 3095: 3093: 3091: 3058: 3052: 3051: 3041: 3031: 2999: 2993: 2992: 2990: 2988: 2967: 2958: 2952: 2951: 2949: 2947: 2931: 2925: 2924: 2922: 2920: 2905: 2899: 2898: 2888: 2878: 2846: 2840: 2839: 2837: 2835: 2830:on 27 April 2011 2815: 2809: 2808: 2777: 2771: 2770: 2752: 2746: 2745: 2735: 2696:The ISME Journal 2688:Girguis, Peter R 2683: 2677: 2676: 2666: 2640: 2631:(6093): 470–73. 2616: 2610: 2609: 2591: 2574:(6093): 467–70. 2555: 2549: 2548: 2546: 2544: 2529: 2523: 2522: 2496: 2472: 2466: 2465: 2455: 2431: 2425: 2424: 2414: 2394: 2388: 2387: 2377: 2367: 2335: 2329: 2328: 2292: 2269: 2268: 2256: 2250: 2249: 2247: 2245: 2239: 2228: 2220: 2214: 2213: 2203: 2193: 2169: 2163: 2162: 2160: 2158: 2143: 2137: 2136: 2134: 2132: 2118: 2112: 2111: 2085: 2044: 2035: 2034: 2031:10.1038/ngeo1773 2004: 1998: 1997: 1995: 1993: 1978: 1967: 1966: 1964: 1962: 1956: 1951:. Archived from 1938: 1932: 1931: 1929: 1927: 1915: 1909: 1908: 1906: 1904: 1898: 1890: 1884: 1878: 1873: 1863: 1853: 1829: 1823: 1822: 1812: 1789:Current Genomics 1780: 1774: 1773: 1754:10.1038/35059215 1726:Rothschild, Lynn 1722: 1713: 1712: 1694: 1684: 1660: 933:Escherichia coli 773:polyextremophile 768:Polyextremophile 678:Hyperthermophile 461:Halobacteriaceae 327:Natronobacterium 242:High temperature 220: 219: 175:bacterial spores 86: 79: 21: 6002: 6001: 5997: 5996: 5995: 5993: 5992: 5991: 5982:Geomicrobiology 5947: 5946: 5945: 5933: 5923: 5921: 5913: 5911: 5906: 5897:Thermostability 5833:Grylloblattidae 5803:Acidobacteriota 5777: 5731: 5692: 5626: 5620: 5562:Metallotolerant 5500: 5495: 5449:Wayback Machine 5427:Wayback Machine 5410:Wayback Machine 5394: 5389: 5383: 5281: 5279:Further reading 5276: 5275: 5220: 5216: 5177: 5173: 5134: 5127: 5088: 5084: 5042: 5036: 5027: 4990: 4983: 4944: 4940: 4895: 4891: 4836: 4832: 4795: 4791: 4784: 4766: 4762: 4752: 4750: 4737: 4736: 4732: 4724: 4720: 4714:Wayback Machine 4705: 4701: 4661: 4657: 4618: 4614: 4556: 4552: 4541: 4537: 4494: 4490: 4477: 4476: 4472: 4453: 4449: 4394: 4390: 4351: 4347: 4341: 4310: 4309: 4305: 4260: 4256: 4195: 4191: 4136: 4132: 4122: 4120: 4112: 4111: 4107: 4097: 4095: 4085: 4081: 4065: 4064: 4057: 4055: 4045: 4041: 4032: 4030: 4020: 3996: 3992: 3949: 3945: 3897: 3893: 3860:(12): 2239–51. 3846: 3842: 3813:(1–3): 117–25. 3803: 3799: 3784: 3780: 3765: 3761: 3708: 3704: 3646: 3642: 3605:(5–6): 650–63. 3595: 3591: 3532: 3528: 3521: 3499: 3495: 3428: 3424: 3414: 3412: 3400: 3396: 3323: 3319: 3309: 3307: 3298: 3297: 3293: 3283: 3281: 3273: 3272: 3268: 3237: 3233: 3180: 3176: 3166: 3164: 3152: 3148: 3132:10.1038/512244a 3103: 3099: 3089: 3087: 3059: 3055: 3000: 2996: 2986: 2984: 2965: 2959: 2955: 2945: 2943: 2932: 2928: 2918: 2916: 2914:Harvard Gazette 2906: 2902: 2847: 2843: 2833: 2831: 2816: 2812: 2805: 2778: 2774: 2767: 2753: 2749: 2684: 2680: 2617: 2613: 2556: 2552: 2542: 2540: 2531: 2530: 2526: 2494:10.1038/nbt1010 2487:(10): 1275–81. 2473: 2469: 2432: 2428: 2395: 2391: 2336: 2332: 2293: 2272: 2257: 2253: 2243: 2241: 2237: 2226: 2222: 2221: 2217: 2170: 2166: 2156: 2154: 2144: 2140: 2130: 2128: 2120: 2119: 2115: 2045: 2038: 2005: 2001: 1991: 1989: 1979: 1970: 1960: 1958: 1939: 1935: 1925: 1923: 1916: 1912: 1902: 1900: 1896: 1892: 1891: 1887: 1830: 1826: 1781: 1777: 1723: 1716: 1661: 1654: 1649: 1600: 1592:Methanococcales 1483: 1430: 1424: 1386: 1374:gamma radiation 1327: 1321: 1309: 1273: 1205: 1148: 1135: 988:simulation time 944:ultracentrifuge 853: 851:In astrobiology 688:Metallotolerant 664:Hyperpiezophile 569: 529: 527:Classifications 454: 376:1,500 to 6,000 272:Low temperature 211:protein folding 191:Marianas Trench 154: 143: 141:Characteristics 28: 23: 22: 15: 12: 11: 5: 6000: 5990: 5989: 5984: 5979: 5974: 5969: 5964: 5959: 5944: 5943: 5931: 5908: 5907: 5905: 5904: 5899: 5894: 5886: 5881: 5876: 5871: 5866: 5859: 5854: 5847: 5840: 5835: 5830: 5828:Thermoproteota 5825: 5820: 5815: 5810: 5805: 5800: 5795: 5789: 5787: 5783: 5782: 5779: 5778: 5776: 5775: 5770: 5763: 5756: 5749: 5741: 5739: 5733: 5732: 5730: 5729: 5722: 5715: 5710: 5702: 5700: 5694: 5693: 5691: 5690: 5685: 5678: 5671: 5664: 5659: 5654: 5647: 5639: 5637: 5628: 5622: 5621: 5619: 5618: 5613: 5608: 5599: 5597:Radioresistant 5594: 5589: 5584: 5579: 5574: 5569: 5564: 5559: 5554: 5549: 5547:Lithoautotroph 5544: 5539: 5534: 5529: 5524: 5519: 5514: 5508: 5506: 5502: 5501: 5494: 5493: 5486: 5479: 5471: 5465: 5464: 5459: 5451: 5439: 5434: 5429: 5417: 5412: 5400: 5393: 5392:External links 5390: 5388: 5387: 5381: 5368: 5360:"Extremophile" 5355: 5312: 5282: 5280: 5277: 5274: 5273: 5214: 5171: 5144:(4): 1167–75. 5125: 5082: 5025: 4981: 4938: 4889: 4830: 4809:(10): 767–78. 4789: 4782: 4760: 4730: 4718: 4699: 4672:(10): 859–72. 4655: 4612: 4550: 4548:15 April 2010. 4535: 4508:(4): 301–315. 4488: 4470: 4447: 4388: 4345: 4339: 4303: 4274:(3): 450–451. 4254: 4189: 4130: 4105: 4079: 4039: 4018: 3990: 3943: 3891: 3840: 3797: 3778: 3759: 3722:(9): 1981–86. 3702: 3640: 3589: 3526: 3519: 3493: 3422: 3394: 3317: 3291: 3266: 3231: 3174: 3146: 3097: 3053: 2994: 2953: 2926: 2900: 2841: 2810: 2803: 2772: 2766:978-0805075120 2765: 2747: 2702:(7): 1545–58. 2678: 2611: 2550: 2524: 2467: 2426: 2389: 2330: 2303:(4): 785–811. 2270: 2251: 2215: 2164: 2138: 2113: 2036: 1999: 1968: 1933: 1910: 1885: 1824: 1795:(3): 161–167. 1775: 1714: 1651: 1650: 1648: 1645: 1644: 1643: 1638: 1633: 1627: 1621: 1616: 1611: 1606: 1599: 1596: 1588:Thermococcales 1489:The bacterium 1482: 1479: 1426:Main article: 1423: 1420: 1385: 1382: 1320: 1317: 1308: 1305: 1272: 1269: 1204: 1201: 1147: 1144: 1139:bioremediation 1134: 1133:Bioremediation 1131: 852: 849: 848: 847: 844:water activity 840: 834: 833: 830: 824: 823: 810: 806: 805: 794: 792:Radioresistant 788: 787: 784: 777: 776: 769: 765: 764: 753: 747: 746: 743: 737: 736: 733: 727: 726: 690: 684: 683: 680: 674: 673: 666: 660: 659: 656: 650: 649: 639: 633: 632: 625: 619: 618: 611: 568: 565: 546:radioresistant 542:Atacama Desert 528: 525: 522: 521: 507: 505: 502: 498: 497: 490: 487: 480:Atacama Desert 477: 471: 470: 457: 452: 448: 445: 441: 440: 432: 426: 424:Mariana Trench 421: 417: 416: 409: 406: 400: 399: 380: 374: 361: 355: 354: 347: 346:pH 0.06 to 1.0 344: 338: 331: 330: 305: 299: 294: 287: 286: 279: 276: 273: 269: 268: 255: 252: 243: 239: 238: 235: 232: 229: 225: 224: 160:microbial life 149: 142: 139: 135:bioremediation 26: 9: 6: 4: 3: 2: 5999: 5988: 5985: 5983: 5980: 5978: 5975: 5973: 5970: 5968: 5965: 5963: 5960: 5958: 5957:Extremophiles 5955: 5954: 5952: 5942: 5937: 5932: 5930: 5920: 5919: 5916: 5903: 5900: 5898: 5895: 5893: 5891: 5887: 5885: 5882: 5880: 5877: 5875: 5872: 5870: 5867: 5865: 5864: 5860: 5858: 5855: 5853: 5852: 5848: 5846: 5845: 5844:Halobacterium 5841: 5839: 5836: 5834: 5831: 5829: 5826: 5824: 5821: 5819: 5816: 5814: 5811: 5809: 5806: 5804: 5801: 5799: 5796: 5794: 5791: 5790: 5788: 5784: 5774: 5771: 5769: 5768: 5764: 5762: 5761: 5757: 5755: 5754: 5750: 5748: 5747: 5743: 5742: 5740: 5738: 5734: 5728: 5727: 5723: 5721: 5720: 5716: 5714: 5711: 5709: 5708: 5704: 5703: 5701: 5699: 5695: 5689: 5686: 5684: 5683: 5679: 5677: 5676: 5672: 5670: 5669: 5665: 5663: 5660: 5658: 5655: 5653: 5652: 5648: 5646: 5645: 5641: 5640: 5638: 5636: 5632: 5629: 5627:extremophiles 5623: 5617: 5614: 5612: 5609: 5607: 5603: 5600: 5598: 5595: 5593: 5590: 5588: 5585: 5583: 5580: 5578: 5575: 5573: 5570: 5568: 5565: 5563: 5560: 5558: 5555: 5553: 5550: 5548: 5545: 5543: 5540: 5538: 5535: 5533: 5530: 5528: 5525: 5523: 5520: 5518: 5515: 5513: 5510: 5509: 5507: 5503: 5499: 5498:Extremophiles 5492: 5487: 5485: 5480: 5478: 5473: 5472: 5469: 5463: 5460: 5458: 5457: 5452: 5450: 5446: 5443: 5440: 5438: 5435: 5433: 5430: 5428: 5424: 5421: 5418: 5416: 5413: 5411: 5407: 5404: 5401: 5399: 5396: 5395: 5384: 5378: 5374: 5369: 5365: 5361: 5356: 5352: 5348: 5343: 5338: 5334: 5330: 5326: 5322: 5318: 5313: 5309: 5305: 5301: 5297: 5293: 5289: 5284: 5283: 5269: 5265: 5260: 5255: 5250: 5245: 5241: 5237: 5234:(1): e49044. 5233: 5229: 5225: 5218: 5210: 5206: 5202: 5198: 5194: 5190: 5187:(1): 109–16. 5186: 5182: 5175: 5167: 5163: 5159: 5155: 5151: 5147: 5143: 5139: 5132: 5130: 5121: 5117: 5113: 5109: 5105: 5101: 5098:(4): 545–63. 5097: 5093: 5092:Extremophiles 5086: 5078: 5074: 5070: 5066: 5061: 5056: 5053:(4): 807–17. 5052: 5048: 5041: 5034: 5032: 5030: 5021: 5017: 5012: 5007: 5004:(4): 938–52. 5003: 4999: 4995: 4988: 4986: 4977: 4973: 4969: 4965: 4961: 4957: 4953: 4949: 4942: 4934: 4930: 4925: 4920: 4916: 4912: 4909:(1): 338–40. 4908: 4904: 4900: 4893: 4885: 4881: 4876: 4871: 4866: 4861: 4857: 4853: 4850:(1): 176–83. 4849: 4845: 4841: 4834: 4826: 4822: 4817: 4812: 4808: 4804: 4800: 4793: 4785: 4779: 4775: 4771: 4764: 4748: 4744: 4740: 4734: 4727: 4722: 4715: 4711: 4708: 4703: 4695: 4691: 4687: 4683: 4679: 4675: 4671: 4667: 4659: 4651: 4647: 4643: 4639: 4635: 4631: 4628:(2): 217–24. 4627: 4623: 4622:Extremophiles 4616: 4608: 4604: 4600: 4596: 4592: 4588: 4584: 4580: 4575: 4570: 4567:(3): 241–58. 4566: 4562: 4554: 4547: 4544: 4539: 4531: 4527: 4523: 4519: 4515: 4511: 4507: 4503: 4499: 4492: 4484: 4480: 4474: 4466: 4462: 4458: 4451: 4443: 4439: 4434: 4429: 4425: 4421: 4416: 4411: 4407: 4403: 4399: 4392: 4384: 4380: 4376: 4372: 4368: 4364: 4360: 4356: 4349: 4342: 4336: 4331: 4326: 4322: 4318: 4314: 4307: 4298: 4293: 4289: 4285: 4281: 4277: 4273: 4269: 4265: 4258: 4250: 4246: 4241: 4236: 4232: 4228: 4224: 4220: 4216: 4212: 4208: 4204: 4200: 4193: 4185: 4181: 4176: 4171: 4167: 4163: 4158: 4153: 4149: 4145: 4141: 4134: 4119: 4115: 4109: 4094: 4090: 4083: 4075: 4069: 4054: 4050: 4043: 4029: 4025: 4021: 4015: 4011: 4007: 4003: 4002: 3994: 3986: 3982: 3978: 3974: 3970: 3966: 3962: 3958: 3954: 3947: 3939: 3935: 3931: 3927: 3923: 3919: 3915: 3911: 3907: 3903: 3895: 3887: 3883: 3878: 3873: 3868: 3863: 3859: 3855: 3851: 3844: 3836: 3832: 3828: 3824: 3820: 3816: 3812: 3808: 3801: 3793: 3789: 3782: 3774: 3770: 3763: 3755: 3751: 3747: 3743: 3739: 3735: 3730: 3725: 3721: 3717: 3713: 3706: 3698: 3694: 3690: 3686: 3682: 3678: 3673: 3668: 3664: 3660: 3657:(4): 600–19. 3656: 3652: 3644: 3636: 3632: 3628: 3624: 3620: 3616: 3612: 3608: 3604: 3600: 3593: 3585: 3581: 3576: 3571: 3567: 3563: 3558: 3553: 3549: 3545: 3541: 3537: 3530: 3522: 3520:9780198726289 3516: 3512: 3508: 3504: 3497: 3490: 3485: 3479: 3475: 3471: 3467: 3462: 3457: 3452: 3447: 3443: 3439: 3438: 3433: 3426: 3411: 3410: 3405: 3398: 3391: 3386: 3380: 3376: 3372: 3368: 3363: 3358: 3354: 3350: 3345: 3340: 3336: 3332: 3328: 3321: 3305: 3301: 3295: 3280: 3276: 3270: 3262: 3258: 3254: 3250: 3246: 3242: 3235: 3227: 3223: 3218: 3213: 3209: 3205: 3201: 3197: 3194:(3): 231–37. 3193: 3189: 3185: 3178: 3163: 3162: 3157: 3150: 3142: 3138: 3133: 3128: 3124: 3120: 3116: 3112: 3108: 3101: 3086: 3082: 3078: 3074: 3070: 3069: 3064: 3057: 3049: 3045: 3040: 3035: 3030: 3025: 3021: 3017: 3014:(4): e62437. 3013: 3009: 3005: 2998: 2983: 2979: 2975: 2971: 2964: 2957: 2941: 2937: 2930: 2915: 2911: 2904: 2896: 2892: 2887: 2882: 2877: 2872: 2868: 2864: 2860: 2856: 2852: 2845: 2829: 2825: 2821: 2814: 2806: 2800: 2796: 2792: 2788: 2784: 2776: 2768: 2762: 2758: 2751: 2743: 2739: 2734: 2729: 2725: 2721: 2717: 2713: 2709: 2705: 2701: 2697: 2693: 2689: 2682: 2674: 2670: 2665: 2660: 2656: 2652: 2648: 2644: 2639: 2634: 2630: 2626: 2622: 2615: 2607: 2603: 2599: 2595: 2590: 2585: 2581: 2577: 2573: 2569: 2565: 2562:(July 2012). 2561: 2554: 2539:. 9 July 2012 2538: 2534: 2528: 2520: 2516: 2512: 2508: 2504: 2500: 2495: 2490: 2486: 2482: 2478: 2471: 2463: 2459: 2454: 2449: 2446:(2): 611–29. 2445: 2441: 2437: 2430: 2422: 2418: 2413: 2408: 2404: 2400: 2393: 2385: 2381: 2376: 2371: 2366: 2361: 2357: 2353: 2350:(6): e66207. 2349: 2345: 2341: 2334: 2326: 2322: 2318: 2314: 2310: 2306: 2302: 2298: 2291: 2289: 2287: 2285: 2283: 2281: 2279: 2277: 2275: 2266: 2262: 2255: 2236: 2232: 2225: 2219: 2211: 2207: 2202: 2197: 2192: 2187: 2183: 2179: 2175: 2168: 2153: 2149: 2142: 2127: 2123: 2117: 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