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''Pyrococcus furiosus'' is a
heterotrophic A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
, strictly
anaerobic Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to: * Anaerobic adhesive, a bonding a ...
, extremophilic, model species of
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaeba ...
. It is classified as a hyperthermophile because it thrives best under extremely high temperatures, and is notable for having an optimum growth temperature of 100 °C (a temperature that would destroy most living organisms). ''P. furiosus'' belongs to the ''
Pyrococcus ''Pyrococcus'' is a genus of Thermococcaceaen archaean.See the NCBIbr>webpage on Pyrococcus Data extracted from the Description and significance ''Pyrococcus'' has similar characteristics of other thermoautotrophican archaea such as ''Ar ...
'' genus, most commonly found in extreme environmental conditions of
hydrothermal vents A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspo ...
. It is one of the few prokaryotic organisms that has
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. ...
containing
tungsten Tungsten, or wolfram, is a chemical element with the symbol W and atomic number 74. Tungsten is a rare metal found naturally on Earth almost exclusively as compounds with other elements. It was identified as a new element in 1781 and first isol ...
, an element rarely found in biological molecules. ''Pyrococcus furiosus'' has many potential industrial applications, owing to its unique thermostable properties. ''P. furiosus'' is used in the process of DNA amplification by way of
polymerase chain reaction The polymerase chain reaction (PCR) is a method widely used to rapidly make millions to billions of copies (complete or partial) of a specific DNA sample, allowing scientists to take a very small sample of DNA and amplify it (or a part of it) ...
(PCR) because of its proofreading activity. Utilizing ''P. furiosus'' in PCR DNA amplification instead of the traditionally used ''Taq'' DNA polymerase allows for a significantly more accurate process. The thermodynamic stability of ''P. furiosus enzymes is useful in the creation of diols for laboratory and industrial purposes. Certain
superoxide dismutase Superoxide dismutase (SOD, ) is an enzyme that alternately catalyzes the dismutation (or partitioning) of the superoxide () radical into ordinary molecular oxygen (O2) and hydrogen peroxide (). Superoxide is produced as a by-product of oxygen ...
s found in ''P. furiosus'' can be introduced into plants to increase their tolerance in environmentally stressful conditions and ultimately their survival.


Properties

''Pyrococcus furiosus'' is a strictly anaerobic,
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic, sulfur-reducing archaea originally isolated from heated sediments in Vulcano, Italy by Fiala and Stetter. It is noted for its rapid doubling time of 37 minutes under optimal conditions, meaning that every 37 minutes the number of individual organisms is multiplied by two, yielding an exponential growth curve. Each organism is surrounded by a cellular envelope composed of
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glyco ...
called an S-layer. It appears as mostly regular cocci—meaning that it is roughly spherical—of 0.8 µm to 1.5 µm diameter with monopolar polytrichous
flagellation Flagellation (Latin , 'whip'), flogging or whipping is the act of beating the human body with special implements such as whips, rods, switches, the cat o' nine tails, the sjambok, the knout, etc. Typically, flogging has been imposed on ...
. A
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glyco ...
notable to archaea species makes up the majority of the composition of ''P. furiosus''
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have fro ...
. Aside from potentially using them for swimming, these flagella were observed under lab conditions in use for unique applications such as forming cell to cell connections during stationary growth phase. They are additionally utilized as cable-like connectors to adhere to various solid surfaces such as sand grains in the habitat in which this species was discovered. This may lead to the formation of microcolonial
biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular po ...
-like structures. ''P.'' ''furiosus'' grows between 70
°C The degree Celsius is the unit of temperature on the Celsius scale (originally known as the centigrade scale outside Sweden), one of two temperature scales used in the International System of Units (SI), the other being the Kelvin scale. The d ...
(158 °F) and 103 °C (217 °F), with an optimum temperature of 100 °C (212 °F), and between pH 5 and 9 (with an optimum at pH 7). Since it uses fermentation of carbohydrates, it grows well on yeast extract,
maltose } Maltose ( or ), also known as maltobiose or malt sugar, is a disaccharide formed from two units of glucose joined with an α(1→4) bond. In the isomer isomaltose, the two glucose molecules are joined with an α(1→6) bond. Maltose is the tw ...
,
cellobiose Cellobiose is a disaccharide with the formula (C6H7(OH)4O)2O. It is classified as a reducing sugar. In terms of its chemical structure, it is derived from the condensation of a pair of β-glucose molecules forming a β(1→4) bond. It can be h ...
, β-glucans,
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human die ...
, and protein sources (tryptone, peptone, casein, and meat extracts) through the Embden-Meyerhoff pathway. This is a relatively wide range of sources when compared to other archaea. Growth is very slow, or nonexistent, on amino acids, organic acids, alcohols, and most carbohydrates (including
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...
,
fructose Fructose, or fruit sugar, is a ketonic simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorb ...
,
lactose Lactose is a disaccharide sugar synthesized by galactose and glucose subunits and has the molecular formula C12H22O11. Lactose makes up around 2–8% of milk (by mass). The name comes from ' (gen. '), the Latin word for milk, plus the suffix ' ...
, and
galactose Galactose (, '' galacto-'' + ''-ose'', "milk sugar"), sometimes abbreviated Gal, is a monosaccharide sugar that is about as sweet as glucose, and about 65% as sweet as sucrose. It is an aldohexose and a C-4 epimer of glucose. A galactose molecu ...
). The metabolic products of ''P. furiosus'' are CO2 and H2. The presence of hydrogen severely inhibits its growth and metabolism; this effect can be circumvented, however, by introducing
sulfur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formul ...
into the organism's environment. In this case, H2S can be produced through its metabolic processes seemingly for the purpose of detoxication or energy conservation, not energy production. While many other hyperthermophiles depend on sulfur for growth, ''P. furiosus'' does not. ''P. furiosus'' is also notable for an unusual and intriguingly simple respiratory system, which obtains energy by reducing protons to hydrogen gas and uses this energy to create an electrochemical gradient across its cell membrane, thereby driving
ATP synthesis ATP synthase is a protein that catalyzes the formation of the energy storage molecule adenosine triphosphate (ATP) using adenosine diphosphate (ADP) and inorganic phosphate (Pi). It is classified under ligases as it changes ADP by the formation ...
. This could be a very early evolutionary precursor of respiratory systems in all higher organisms today.


Genomics

The sequencing of the complete
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
of ''Pyrococcus furiosus'' was completed in 2001 by scientists at the University of Maryland Biotechnology Institute. The Maryland team found that the genome has 1,908 kilobases, including 2,065 open reading frames (ORFs) that encode proteins. A study performed in 2005 revealed 17 new ORFs specific to ''Pyrococcus furiosus'' that were not originally annotated, bringing the number of ORFs up to 2,082. A lab strain of ''Pyrococcus furiosus'' named COM1 is commonly used for its "high plasticity" and ability to take up foreign DNA, owing to its high recombination and transposon activity. It has 1,571 more base pairs than the referenced NCBI genome, and 10 more
insertion sequences Insertion element (also known as an IS, an insertion sequence element, or an IS element) is a short DNA sequence that acts as a simple transposable element. Insertion sequences have two major characteristics: they are small relative to other transp ...
(ISs). These ISs have deactivated 13 genes and many more are altered, but the strain's growth is yet comparable to its parent strain.


Enzymes


Alcohol dehydrogenases

''Pyrococcus furiosus'' possesses several highly thermostable alcohol dehydrogenases (ADHs): the short-chain AdhA, the iron-containing AdhB, the zinc-containing AdhC, and more. Each of these ADHs are NADP-dependent, and serve to replenish NADP+ by using the NADPH produced by
fermentation Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food p ...
to reduce aldehydes to alcohols. The aldehydes are also products of fermentation and are toxic to the cell, so removal is necessary. ''P. furiosus'' ADHs typically have a broad range of aldehyde substrates they can use, and they can also catalyze the reverse reaction (oxidation of alcohols) using ethanol, 1,3-propanediol, and other alcohols for substrate. As with most of the archaea's enzymes, the ADHs are sensitive to oxygen.


Oxidoreductases

''Pyrococcus furiosus'' has five unique tungsten-containing
oxidoreductase In biochemistry, an oxidoreductase is an enzyme that catalyzes the transfer of electrons from one molecule, the reductant, also called the electron donor, to another, the oxidant, also called the electron acceptor. This group of enzymes usually ...
s that are part of its NAD(P)H-independent glycolytic pathway. These enzymes function optimally above 90 °C. The first to be discovered was
aldehyde In organic chemistry, an aldehyde () is an organic compound containing a functional group with the structure . The functional group itself (without the "R" side chain) can be referred to as an aldehyde but can also be classified as a formyl gro ...
ferredoxin Ferredoxins (from Latin ''ferrum'': iron + redox, often abbreviated "fd") are iron–sulfur proteins that mediate electron transfer in a range of metabolic reactions. The term "ferredoxin" was coined by D.C. Wharton of the DuPont Co. and applied t ...
oxidoreductase, or AOR, which utilizes tungsten, sulfur, and iron to catalyze the
oxidation Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or ...
of aldehydes and reduce ferredoxin (this being the electron carrier instead of NAD(P)H). As this was the first, all tungsten-containing oxidoreductases are said to be part of the AOR family. The next oxidoreductase to be discovered was
glyceraldehyde-3-phosphate Glyceraldehyde 3-phosphate, also known as triose phosphate or 3-phosphoglyceraldehyde and abbreviated as G3P, GA3P, GADP, GAP, TP, GALP or PGAL, is a metabolite that occurs as an intermediate in several central pathways of all organisms.Nelson, D ...
ferredoxin oxidoreductase, or GAPOR, which utilizes tungsten and iron to catalyze the oxidation of specifically glyceraldehyde-3-phosphate. GAPOR only functions under anaerobic conditions, as with many enzymes in ''P. furiosus''. Another oxidoreductase is
formaldehyde Formaldehyde ( , ) ( systematic name methanal) is a naturally occurring organic compound with the formula and structure . The pure compound is a pungent, colourless gas that polymerises spontaneously into paraformaldehyde (refer to section ...
ferredoxin oxidoreductase, or FOR, which catalyzes the oxidation of aldehydes into
carboxylic acid In organic chemistry, a carboxylic acid is an organic acid that contains a carboxyl group () attached to an R-group. The general formula of a carboxylic acid is or , with R referring to the alkyl, alkenyl, aryl, or other group. Carboxyli ...
s. This enzyme utilizes four types of cofactors: tungsten, iron, sulfur, and calcium. The next oxidoreductase, WOR4, does not help oxidize aldehydes, but rather has a role in the reduction of elemental sulfur (S0) into H2S. This uses the same cofactors as FOR, and is only found in P. furiosus cells that are grown in the presence of elemental sulfur. The fifth and final oxidoreductase is named WOR5, and it has a broad specificity for
aromatic In chemistry, aromaticity is a chemical property of cyclic ( ring-shaped), ''typically'' planar (flat) molecular structures with pi bonds in resonance (those containing delocalized electrons) that gives increased stability compared to satur ...
and
aliphatic In organic chemistry, hydrocarbons ( compounds composed solely of carbon and hydrogen) are divided into two classes: aromatic compounds and aliphatic compounds (; G. ''aleiphar'', fat, oil). Aliphatic compounds can be saturated, like hexane ...
aldehyde species. An oxidoreductase species in ''P. furiosus'' that does not contain tungsten is
pyruvate Pyruvic acid (CH3COCOOH) is the simplest of the alpha-keto acids, with a carboxylic acid and a ketone functional group. Pyruvate, the conjugate base, CH3COCOO−, is an intermediate in several metabolic pathways throughout the cell. Pyruvic a ...
ferredoxin oxidoreductase, or POR, which catalyzes the final step of the glycolytic pathway. It is possible that POR is an ancestor of mesophilic pyruvate oxidoreductases. There is also the indolepyruvate ferredoxin oxidoreductase, or IOR, which utilizes iron and sulfur to catalyze the "oxidative
decarboxylation Decarboxylation is a chemical reaction that removes a carboxyl group and releases carbon dioxide (CO2). Usually, decarboxylation refers to a reaction of carboxylic acids, removing a carbon atom from a carbon chain. The reverse process, which is ...
of
aryl In organic chemistry, an aryl is any functional group or substituent derived from an aromatic ring, usually an aromatic hydrocarbon, such as phenyl and naphthyl. "Aryl" is used for the sake of abbreviation or generalization, and "Ar" is used ...
pyruvates."


Uses


In DNA amplification

A
DNA polymerase A DNA polymerase is a member of a family of enzymes that catalyze the synthesis of DNA molecules from nucleoside triphosphates, the molecular precursors of DNA. These enzymes are essential for DNA replication and usually work in groups to crea ...
was discovered in ''P. furiosus'' that was thought to be unrelated to other known DNA polymerases, as no significant
sequence homology Sequence homology is the biological homology between DNA, RNA, or protein sequences, defined in terms of shared ancestry in the evolutionary history of life. Two segments of DNA can have shared ancestry because of three phenomena: either a ...
was found between its two proteins and those of other known DNA polymerases. This DNA polymerase has strong 3'-to-5' exonucleolytic activity and a template-primer preference which is characteristic of a replicative DNA polymerase, leading scientists to believe that this enzyme may be the replicative DNA polymerase of ''P. furiosus''. It has since been placed in the family B of polymerases, the same family as DNA polymerase II. Its structure, which appears quite typical for polymerase B, has been solved as well. Since the
enzymes Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. ...
of ''P. furiosus'' are extremely thermostable, the
DNA polymerase A DNA polymerase is a member of a family of enzymes that catalyze the synthesis of DNA molecules from nucleoside triphosphates, the molecular precursors of DNA. These enzymes are essential for DNA replication and usually work in groups to crea ...
from ''P. furiosus'' (also known as
Pfu DNA polymerase ''Pfu'' DNA polymerase is an enzyme found in the hyperthermophilic archaeon ''Pyrococcus furiosus'', where it functions to copy the organism's DNA during cell division. In the laboratory setting, ''Pfu'' is used to amplify DNA in the polymeras ...
) can be used in the
polymerase chain reaction The polymerase chain reaction (PCR) is a method widely used to rapidly make millions to billions of copies (complete or partial) of a specific DNA sample, allowing scientists to take a very small sample of DNA and amplify it (or a part of it) ...
(PCR) DNA amplification process. The PCR process must use a thermostable DNA polymerase for automated in vitro amplification and originally used ''Taq'' DNA polymerase. However, since purified ''Taq'' DNA polymerase lacks
exonuclease Exonucleases are enzymes that work by cleaving nucleotides one at a time from the end (exo) of a polynucleotide chain. A hydrolyzing reaction that breaks phosphodiester bonds at either the 3′ or the 5′ end occurs. Its close relative is ...
(proofreading) activity, it cannot excise mismatched
nucleotide Nucleotides are organic molecules consisting of a nucleoside and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecu ...
s. Researchers discovered in the early 1990's that the ''Pfu'' DNA polymerase of ''P. furiosus'' does actually possess a requisite 3’-to-5’ exonuclease activity allowing for the removal of errors. Subsequent tests utilizing ''Pfu'' DNA polymerase in the PCR process revealed a more than tenfold improvement over the accuracy of using ''Taq'' DNA polymerase.


In production of diols

One practical application of ''P. furiosus'' is in the production of
diol A diol is a chemical compound containing two hydroxyl groups ( groups). An aliphatic diol is also called a glycol. This pairing of functional groups is pervasive, and many subcategories have been identified. The most common industrial diol is e ...
s for various industrial processes. It may be possible to use the enzymes of ''P. furiosus'' for applications in such industries as food, pharmaceuticals, and fine-chemicals in which alcohol dehydrogenases are necessary in the production of enantio- and diastereomerically pure diols. Enzymes from hyperthermophiles such as ''P. furiosus'' can perform well in laboratory processes because they are relatively resistant: they generally function well at high temperatures and high pressures, as well as in high concentrations of chemicals. In order to make naturally derived enzymes useful in the laboratory, it is often necessary to alter their genetic makeup. Otherwise, the naturally occurring enzymes may not be efficient in an artificially induced procedure. Although the enzymes of ''P. furiosus'' function optimally at a high temperature, scientists may not necessarily want to carry out a procedure at . Consequently, in this case, the specific enzyme AdhA was taken from ''P. furiosus'' and put through various mutations in a laboratory in order to obtain a suitable alcohol dehydrogenase for use in artificial processes. This allowed scientists to obtain a mutant enzyme that could function efficiently at lower temperatures and maintain productivity.


In plants

The expression of a certain gene found in ''P. furiosus'' in plants can also render them more durable by increasing their tolerance for heat. In response to environmental stresses such as heat exposure, plants produce
reactive oxygen species In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen. The reduction of molecular oxygen () p ...
which can result in cell death. If these free radicals are removed, cell death can be delayed. Enzymes in plants called
superoxide dismutase Superoxide dismutase (SOD, ) is an enzyme that alternately catalyzes the dismutation (or partitioning) of the superoxide () radical into ordinary molecular oxygen (O2) and hydrogen peroxide (). Superoxide is produced as a by-product of oxygen ...
s remove superoxide anion radicals from cells, but increasing the amount and activity of these enzymes is difficult and not the most efficient way to go about improving the durability of plants. By introducing the
superoxide reductase Superoxide reductase is an enzyme that catalyzes the conversion of highly reactive and toxic superoxide (O2−) into less toxic hydrogen peroxide (H2O2): :reduced rubredoxin + O2− + 2 H+ \rightleftharpoons rubredoxin + H2O2 :Fe2+ + O2− + 2 H+ ...
s of ''P. furiosus'' into plants, the levels of O2 can be rapidly reduced. Scientists tested this method using the ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter ...
'' plant. As a result of this procedure, cell death in plants occurs less often, therefore resulting in a reduction in the severity of responses to environmental stress. This enhances the survival of plants, making them more resistant to light, chemical, and heat stress. This study could potentially be used as a starting point to creating plants that could survive in more extreme climates on other planets such as Mars. By introducing more enzymes from extremophiles like ''P. furiosus'' into other species of plants, it may be possible to create incredibly resistant species.


In researching amino acids

By comparing ''P. furiosus'' with a related species of archaea, ''
Pyrococcus abyssi ''Pyrococcus abyssi'' is a hyperthermophilic archaeon isolated from a deep-sea hydrothermal vent in the North Fiji Basin at . It is anaerobic, sulfur-metabolizing, gram-negative, coccus-shaped and highly motile. Its optimum growth temperature ...
'', scientists have tried to determine the correlation between certain amino acids and affinity for certain pressures in different species. ''P. furiosus'' is not barophilic, while ''P. abyssi'' is, meaning that it functions optimally at very high pressures. Using two hyperthermophilic species of archaea lessens the possibility of deviations having to do with temperature of the environment, essentially reducing the variables in the experimental design. Besides yielding information about the barophilicity of certain amino acids, the experiment also provided valuable insight into the origin of the genetic code and its organizational influences. It was found that most of the amino acids that determined barophilicity were also found to be important in the organization of the genetic code. It was also found that more polar amino acids and smaller amino acids were more likely to be barophilic. Through the comparison of these two archaea, the conclusion was reached that the genetic code was likely structured under high hydrostatic pressure, and that hydrostatic pressure was a more influential factor in determining genetic code than temperature.


History

''Pyrococcus furiosus'' was originally isolated anaerobically from geothermal marine sediments with temperatures between and collected at the beach of Porto Levante,
Vulcano Island Vulcano ( scn, Vurcanu) or Vulcan is a small volcanic island belonging to Italy in the Tyrrhenian Sea, about north of Sicily and located at the southernmost end of the seven Aeolian Islands. The island is known for its volcanic activity and ...
, Italy. It was first described by
Karl Stetter Karl Otto Stetter (born 16 July 1941) is a German microbiologist and authority on astrobiology. He is an expert on microbial life at high temperatures. Career Stetter was born in Munich and studied biology at the Technical University of Munich. ...
of the
University of Regensburg The University of Regensburg (german: link=no, Universität Regensburg) is a public research university located in the medieval city of Regensburg, Bavaria, a city that is listed as a UNESCO World Heritage Site. The university was founded on 18 ...
in Germany, and a colleague, Gerhard Fiala. ''Pyrococcus furiosus'' actually originated a new genus of archaea with its relatively recent discovery in 1986. The name ' means "fireball" in
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, to refer to the extremophile's round shape and ability to grow in temperatures of around 100 degrees Celsius. The species name ' means 'rushing' in
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
, and refers to the extremophile's doubling time and rapid swimming.


References


Further reading

* * * *


External links


Type strain of ''Pyrococcus furiosus'' at Bac''Dive'' - the Bacterial Diversity MetadatabaseKEGG Genome : ''Pyrococcus furiosus''
{{Taxonbar, from=Q146310 Euryarchaeota Archaea described in 1993