An extremophile (from Latin extremus meaning "extreme" and Greek phili? () meaning "love") is an organism with optimal growth in environmental conditions considered extreme in that it is challenging for a carbon-based life form, such as all life on Earth, to survive.
These organisms are dominants in the evolutionary history of the planet. Dating back to more than 40 million years ago, extremophiles have continued to thrive in the most extreme conditions naming them one of the most abundant lifeforms. 
This is not the same as a more anthropocentric and non-scientific view which considers an extremophile to be an organism that lives in environments uncomfortable to humans. In contrast, organisms that live in more moderate environmental conditions, according to an anthropocentric view, may be termed mesophiles or neutrophiles.
In the 1980s and 1990s, biologists found that microbial life has great flexibility for surviving in extreme environments--niches that are acidic, extraordinarily hot or within irregular air pressure for example--that would be completely inhospitable to complex organisms. Some scientists even concluded that life may have begun on Earth in hydrothermal vents far under the ocean's surface.
According to astrophysicist Steinn Sigurdsson, "There are viable bacterial spores that have been found that are 40 million years old on Earth--and we know they're very hardened to radiation." Some bacteria were found living in the cold and dark in a lake buried a half-mile deep under the ice in Antarctica, and in the Marianas Trench, the deepest place in Earth's oceans. 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 adaptable to conditions, and survive wherever they are." A key to extremophile adaptation is their amino acid composition, affecting their protein folding ability under particular conditions. Studying extreme environments on Earth can help researchers understand the limits of habitability on other worlds.
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).
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 polyextremophiles. For example, organisms living inside hot rocks deep under Earth's surface are thermophilic and piezophilic such as Thermococcus barophilus. A polyextremophile living at the summit of a mountain in the Atacama Desert might be a radioresistantxerophile, a psychrophile, and an oligotroph. Polyextremophiles are well known for their ability to tolerate both high and low pH levels.
An organism with optimal growth in the absence of molecular oxygen. Two sub-types exist: facultative anaerobe and obligate anaerobe. A facultative anaerobe can tolerate anoxic and oxic conditions whilst an obligate anaerobe will die in the presence of even low levels of molecular oxygen.
An organism that lives in microscopic spaces within rocks, such as pores between aggregate grains. These may also be called endolith, a term that also includes organisms populating fissures, aquifers, and faults filled with groundwater in the deep subsurface.
Recent research carried out on extremophiles in Japan involved a variety of bacteria including Escherichia coli and Paracoccus denitrificans being subject to conditions of extreme gravity. The bacteria were cultivated while being rotated in an ultracentrifuge at high speeds corresponding to 403,627 g (i.e. 403,627 times the gravity experienced on Earth). Paracoccus denitrificans 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 supernovas. Analysis showed that the small size of prokaryotic cells is essential for successful growth under hypergravity. The research has implications on the feasibility of panspermia.
In September 2015, scientists from CNR-National Research Council of Italy reported that S.soflataricus 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.
In June 2016, scientists from Brigham Young University conclusively reported that endospores of Bacillus subtilis 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 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.
In August 2020 scientists reported that bacteria that feed on air discovered 2017 in Antarctica 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 carbon sink and further strengthens the extremophile evidence that supports the potential existence of microbial life on alien planets.
New sub-types of -philes are identified frequently and the sub-category list for extremophiles is always growing. For example, microbial life lives in the liquid asphalt lake, Pitch Lake. Research indicates that extremophiles inhabit the asphalt lake in populations ranging between 106 to 107 cells/gram. Likewise, until recently boron tolerance was unknown but a strong borophile was discovered in bacteria. With the recent isolation of Bacillus boroniphilus, borophiles came into discussion. Studying these borophiles may help illuminate the mechanisms of both boron toxicity and boron deficiency.
In July 2019, a scientific study of Kidd Mine in Canada discovered sulfur-breathing organisms which live 7900 feet 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.
The thermoalkaliphilic catalase, which initiates the breakdown of hydrogen peroxide into oxygen and water, was isolated from an organism, Thermus brockianus, found in Yellowstone National Park by Idaho National Laboratory 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 T. brockianus catalase exhibited a half life of 15 days at 80 °C and pH 10 while a catalase derived from Aspergillus niger 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.
DNA modifying enzymes such as Taq DNA polymerase and some Bacillus enzymes used in clinical diagnostics and starch liquefaction are produced commercially by several biotechnology companies.
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 transfer, as described below. However, it is not yet clear how common such a capability is among extremophiles.
The bacterium Deinococcus radiodurans 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. D. radiodurans is competent to perform genetic transformation. Recipient cells are able to repair DNA damage in donor transforming DNA that had been UV irradiated as efficiently as they repair cellular DNA when the cells themselves are irradiated. The extreme thermophilic bacterium Thermus thermophilus and other related Thermus species are also capable of genetic transformation.
Halobacterium volcanii, an extreme halophilic (saline 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.
Sulfolobus solfataricus and Sulfolobus acidocaldarius 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 S. acidocaldarius 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 Sulfolobus 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 Sulfolobus 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 Transformation (genetics)).
Extracellular membrane vesicles (MVs) might be involved in DNA transfer between different hyperthermophilic archaeal species. It has been shown that both plasmids and viral genomes can be transferred via MVs. Notably, a horizontal plasmid transfer has been documented between hyperthermophilic Thermococcus and Methanocaldococcus species, respectively belonging to the orders Thermococcales and Methanococcales.
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