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<title>Thermococcus celer</title>
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<span id="openzim-page-title" class="mw-page-title-main"><i>Thermococcus celer</i></span>
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<th colspan="2" style="color:inherit; text-align: center; background-color: rgb(195,245,250)"><i>Thermococcus celer</i>
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<th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(195,245,250)"><a href="Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class=" taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"></span></span>
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<td>Domain:
</td>
<td><a href="Archaea" title="Archaea">Archaea</a>
</td></tr>
<tr class="taxonrow">
<td>Kingdom:
</td>
<td><a href="Methanobacteriati" title="Methanobacteriati">Methanobacteriati</a>
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<tr class="taxonrow">
<td>Phylum:
</td>
<td><a href="Methanobacteriota" title="Methanobacteriota">Methanobacteriota</a>
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<tr class="taxonrow">
<td>Class:
</td>
<td><a href="Thermococci" title="Thermococci">Thermococci</a>
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<td>Order:
</td>
<td><a href="Thermococcales" title="Thermococcales">Thermococcales</a>
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<tr class="taxonrow">
<td>Family:
</td>
<td><a href="Thermococcaceae" title="Thermococcaceae">Thermococcaceae</a>
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<tr class="taxonrow">
<td>Genus:
</td>
<td><a href="Thermococcus" title="Thermococcus"><i>Thermococcus</i></a>
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<tr class="taxonrow">
<td>Species:
</td>
<td><div style="display:inline" class="species"><i><b>T.&nbsp;celer</b></i></div>
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<th colspan="2" style="color:inherit; text-align: center; background-color: rgb(195,245,250)"><a href="Binomial_nomenclature" title="Binomial nomenclature">Binomial name</a>
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<td colspan="2" style="text-align: center"><b><span class="binomial"><i>Thermococcus celer</i></span></b><br><div style="font-size: 85%;">Zillig 1983</div>
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<p><i><b>Thermococcus celer</b></i> is a <a href="Gram-negative" class="mw-redirect" title="Gram-negative">Gram-negative</a>, spherical-shaped <a href="Archaeon" class="mw-redirect" title="Archaeon">archaeon</a> of the genus <i><a href="Thermococcus" title="Thermococcus">Thermococcus</a></i>.<sup id="cite_ref-Achenbach_1-0" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The discovery of <i>T. celer</i> played an important role in rerooting the <a href="Tree_of_life" title="Tree of life">tree of life</a> when <i>T. celer</i> was found to be more closely related to <a href="Methanogenic" class="mw-redirect" title="Methanogenic">methanogenic</a> <a href="Archaea" title="Archaea">Archaea</a> than to other phenotypically similar <a href="Thermophilic" class="mw-redirect" title="Thermophilic">thermophilic</a> species.<sup id="cite_ref-Achenbach_1-1" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <i>T. celer</i> was the first archaeon discovered to house a circularized <a href="Genome" title="Genome">genome</a>.<sup id="cite_ref-Noll_2-0" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Several <a href="Strain_(biology)" title="Strain (biology)">type strains</a> of <i>T. celer</i> have been identified: Vu13, ATCC 35543, and DSM 2476.<sup id="cite_ref-Noll_2-1" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
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<div class="mw-heading mw-heading2"><h2 id="Isolation">Isolation</h2></div>
<p><i>T. celer</i> was discovered by Dr. Wolfram Zillig in 1983.<sup id="cite_ref-Zillig_3-0" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The organism was isolated on the beaches of <a href="Vulcano" title="Vulcano">Vulcano</a>, <a href="Italy" title="Italy">Italy</a>, from a <a href="Sulfur" title="Sulfur">sulfur</a>-rich shallow volcanic crater.<sup id="cite_ref-Zillig_3-1" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
Original samples were isolated from the depths of the marine holes and inoculated into 10-ml anaerobic tubes.<sup id="cite_ref-Stetter_4-0" class="reference"><a href="#cite_note-Stetter-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The tubes contained 100&nbsp;mg of elemental <a href="Sulfur" title="Sulfur">sulfur</a> and a solution of 95% N<sub>2</sub> and 5% H<sub>2</sub>S.<sup id="cite_ref-Stetter_4-1" class="reference"><a href="#cite_note-Stetter-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The <a href="PH" title="PH">pH</a> was subsequently adjusted to a range of 5-6 through the addition of CaCO<sub>3</sub>.<sup id="cite_ref-Stetter_4-2" class="reference"><a href="#cite_note-Stetter-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> To ensure that no <a href="Oxygen" title="Oxygen">oxygen</a> had permeated the sample, researchers used the oxygen indicator <a href="Resazurin" title="Resazurin">resazurin</a>.<sup id="cite_ref-Stetter_4-3" class="reference"><a href="#cite_note-Stetter-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Growth was achieved by enrichment with Brock's <i>Sulfolobus</i> <a href="Growth_medium" title="Growth medium">medium</a>, which contains elemental <a href="Sulfur" title="Sulfur">sulfur</a> and <a href="Yeast" title="Yeast">yeast</a>, both of which are required by <i>T. celer</i> for optimal growth.<sup id="cite_ref-Zillig_3-2" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Following enrichment, the samples were plated onto <a href="Polyacrylamide_gel" class="mw-redirect" title="Polyacrylamide gel">polyacrylamide gel</a> and then incubated at 85&nbsp;°C in an anaerobic environment.<sup id="cite_ref-Stetter_4-4" class="reference"><a href="#cite_note-Stetter-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Once colony growth had been observed, the cells were subjected to <a href="Centrifugation" title="Centrifugation">centrifugation</a> prior to purification in a TA <a href="Buffer_solution" title="Buffer solution">buffer solution</a> (0.05&nbsp;mol/L Tris HCl, 0.022&nbsp;mol/L NH<sub>4</sub>Cl, 0.01&nbsp;mol/L β-mercaptoethanol).<sup id="cite_ref-Stetter_4-5" class="reference"><a href="#cite_note-Stetter-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Taxonomy_and_phylogeny">Taxonomy and phylogeny</h2></div>
<p>Following <a href="Sanger_sequencing" title="Sanger sequencing">Sanger sequencing</a> of the <a href="16s_rRNA" class="mw-redirect" title="16s rRNA">16s rRNA</a>, both <a href="Occam's_razor" title="Occam's razor">parsimony</a> and <a href="Distance_matrix" title="Distance matrix">distance matrix</a> analyses were performed to determine the position of <i>T. celer</i> on the tree of life.<sup id="cite_ref-Achenbach_1-2" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <i>T. celer</i> was found to be more closely related to the <a href="Methanogenic" class="mw-redirect" title="Methanogenic">methanogenic</a> <a href="Archaebacteria" class="mw-redirect" title="Archaebacteria">archaebacteria</a> than the <a href="Thermophilic" class="mw-redirect" title="Thermophilic">thermophilic</a> archaebacteria.<sup id="cite_ref-Achenbach_1-3" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This discovery resulted in a rerooting of the archaebacterial tree and subsequently placed <i>T. celer</i> in a <a href="Clade" title="Clade">clade</a> with the methanogens based upon their close <a href="Phylogenetic" class="mw-redirect" title="Phylogenetic">phylogenetic</a> relationship.<sup id="cite_ref-Achenbach_1-4" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This placement was further supported following analysis of the organizational genome structure of the both species’ <a href="RRNA" class="mw-redirect" title="RRNA">rRNA</a> genes.<sup id="cite_ref-Achenbach_1-5" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Both <i>Thermococcus</i> and methanogenic archaebacteria have a <a href="TRNA" class="mw-redirect" title="TRNA">tRNA</a> spacer gene located between the 16s rRNA gene and 23s[rRNA gene.<sup id="cite_ref-Achenbach_1-6" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This spacer gene is not found in any other thermophilic archaebacteria species.<sup id="cite_ref-Achenbach_1-7" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p><p><i>T. celer</i> is related to <i><a href="Pyrococcus_woesei" title="Pyrococcus woesei">Pyrococcus woesei</a></i>, both belonging to the order <a href="Thermococcales" title="Thermococcales">Thermococcales</a>.<sup id="cite_ref-Blamey_5-0" class="reference"><a href="#cite_note-Blamey-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Both are strictly <a href="Anaerobic_organism" title="Anaerobic organism">anaerobic</a> and <a href="Sulphur" class="mw-redirect" title="Sulphur">sulphur</a>-reducing.<sup id="cite_ref-Blamey_5-1" class="reference"><a href="#cite_note-Blamey-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <i>T. celer</i> also shares a close relationship with <i><a href="Thermococcus_litoralis" title="Thermococcus litoralis">Thermococcus litoralis</a></i>, both belonging to the same <a href="Genus" title="Genus">genus</a>, but <i>T. celer</i> has shown to be much more Sulphur-dependent than <i>T. littorals</i>.<sup id="cite_ref-Blamey_5-2" class="reference"><a href="#cite_note-Blamey-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>
</p><p><i> T. celer</i> is currently classified as a thermophilic <a href="Archaeon" class="mw-redirect" title="Archaeon">Archaeon</a>.<sup id="cite_ref-Zillig_3-3" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Since the discovery of <i>T. celer</i>, the term archaebacteria has been replaced with <a href="Archaea" title="Archaea">Archaea</a> as to reflect the most current phylogenetic relationships discovered between the organisms.<sup id="cite_ref-Pace_6-0" class="reference"><a href="#cite_note-Pace-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Characterization">Characterization</h2></div>
<p><i>Thermococcus</i> is constructed from two <a href="Greek_language" title="Greek language">Greek</a> nouns: <i>therme</i> (<a href="Greek_language" title="Greek language">Greek</a>, meaning heat), and <i>kokkos</i> (Greek, meaning <a href="Grain" title="Grain">grain</a> or <a href="Seed" title="Seed">seed</a>).<sup id="cite_ref-Zillig_3-4" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <i>Celer</i> is derived from the Greek, meaning fast, in reference to the species' high growth rates.<sup id="cite_ref-Zillig_3-5" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Morphology">Morphology</h3></div>
<p><i>T. celer</i> is a <a href="Gram-negative" class="mw-redirect" title="Gram-negative">Gram-negative</a>, spherical organism around 1 μm diameter.<sup id="cite_ref-Zillig_3-6" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Observation using <a href="Electron_microscopy" class="mw-redirect" title="Electron microscopy">electron microscopy</a> revealed that <i> T. celer </i> uses a monopolar polytrichous <a href="Flagellum" title="Flagellum">flagellum</a> for movement.<sup id="cite_ref-Zillig_3-7" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> During replication, <i>T. celer</i> is condensed to a diploform shape as seen by <a href="Phase_contrast_microscopy" class="mw-redirect" title="Phase contrast microscopy">phase contrast microscopy</a>.<sup id="cite_ref-Zillig_3-8" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
</p><p>The <i>T. celer</i> <a href="Plasma_membrane" class="mw-redirect" title="Plasma membrane">plasma membrane</a> possesses large amounts of <a href="Glycerol" title="Glycerol">glycerol</a> <a href="Diether" title="Diether">diether</a> <a href="Lipids" class="mw-redirect" title="Lipids">lipids</a> compared to relatively small amounts of diglycerol tetraether lipids.<sup id="cite_ref-Boone_7-0" class="reference"><a href="#cite_note-Boone-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>
Within glycerol diether lipids, <a href="Phytanyl" class="mw-redirect" title="Phytanyl">phytanyl</a> (C<sub>20</sub>) is the <a href="Hydrocarbon" title="Hydrocarbon">hydrocarbon</a> component, and within diglycerol tetraether lipids, biphytanyl (C<sub>40</sub>) is the hydrocarbon component.<sup id="cite_ref-Zillig_3-9" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The <a href="Cell_wall" title="Cell wall">cell wall</a>, or <a href="S-layer" title="S-layer">S-layer</a>, of <i>T. celer</i> functions as protection from <a href="Cell_lysis" class="mw-redirect" title="Cell lysis">cell lysis</a> as a result of changes in osmotic gradients.<sup id="cite_ref-Zillig_3-10" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The envelope S-layer consists of <a href="Glycoprotein" title="Glycoprotein">glycoprotein</a> subunits arranged into a two-dimensional <a href="Paracrystalline" class="mw-redirect" title="Paracrystalline">paracrystalline</a> hexagonal structure.<sup id="cite_ref-Zillig_3-11" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
The <i>T. celer</i> <a href="Cell_envelope" title="Cell envelope">cell envelope</a> lacks <a href="Muramic_acid" title="Muramic acid">muramic acid</a>, indicating <a href="Antibiotic_resistance" class="mw-redirect" title="Antibiotic resistance">resistance</a> to <a href="Penicillin" title="Penicillin">penicillin</a> and <a href="Vancomycin" title="Vancomycin">vancomycin</a>.<sup id="cite_ref-Zillig_3-12" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Metabolism">Metabolism</h3></div>
<p><i>T. celer</i> is a strict <a href="Anaerobe" class="mw-redirect" title="Anaerobe">anaerobe</a> that uses <a href="Organotrophic" class="mw-redirect" title="Organotrophic">organotrophic</a> metabolism<sup id="cite_ref-Zillig_3-13" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> in the form of <a href="Peptides" class="mw-redirect" title="Peptides">peptides</a> (i.e. from <a href="Yeast" title="Yeast">yeast</a> extract, <a href="Peptone" class="mw-redirect" title="Peptone">peptone</a>, or <a href="Tryptone" title="Tryptone">tryptone</a>) and <a href="Proteins" class="mw-redirect" title="Proteins">proteins</a> (i.e. <a href="Casein" title="Casein">casein</a>) as a carbon source which are oxidized to <a href="Carbon_dioxide" title="Carbon dioxide">carbon dioxide</a> via sulphur respiration.<sup id="cite_ref-Zillig_3-14" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <i>T. celer</i> is unable to use <a href="Carbohydrates" class="mw-redirect" title="Carbohydrates">carbohydrates</a> as a carbon source and is considered a sulphur-dependent organism, as it depends upon the reduction of <a href="Sulphur" class="mw-redirect" title="Sulphur">sulphur</a> to <a href="Hydrogen_sulfide" title="Hydrogen sulfide">hydrogen sulfide</a> for optimal growth.<sup id="cite_ref-Blamey_5-3" class="reference"><a href="#cite_note-Blamey-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Though it is less efficient, <i>T. celer</i> is also able to use <a href="Fermentation" title="Fermentation">fermentation</a>.<sup id="cite_ref-Zillig_3-15" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Unlike most <a href="Prokaryotes" class="mw-redirect" title="Prokaryotes">prokaryotes</a>, <i>T. celer</i> is able to perform <a href="Cellular_respiration" title="Cellular respiration">respiration</a> via the Embden–Meyerhof pathway (<a href="Glycolysis" title="Glycolysis">glycolysis</a>), though it uses an alternative route.<sup id="cite_ref-Gadd_8-0" class="reference"><a href="#cite_note-Gadd-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Ecology">Ecology</h3></div>
<p>Characteristic of the <a href="Hyperthermophilic" class="mw-redirect" title="Hyperthermophilic">hyperthermophilic</a> species, <i>T. celer</i> thrives in extremely hot temperatures.<sup id="cite_ref-Achenbach_1-8" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> More specifically, it is found only in sulphur-rich, shallow volcanic craters of Vulcano, Italy.<sup id="cite_ref-Zillig_3-16" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Temperatures in this specific habitat reach up to 90&nbsp;°C <sup id="cite_ref-Zillig_3-17" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The maximum temperature at which <i>T. celer</i> can grow at is 93&nbsp;°C, optimum growth temperature being 88&nbsp;°C.<sup id="cite_ref-Zillig_3-18" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> It grows best at a pH of 5.8, implying that it is mildly <a href="Acidophilic" class="mw-redirect" title="Acidophilic">acidophilic</a>.<sup id="cite_ref-Zillig_3-19" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Optimal growth is also dependent on the presence of a <a href="NaCl" class="mw-redirect" title="NaCl">NaCl</a> concentration of 40 g/L, further demonstrating the high level of adaptation <i>T. celer</i> has evolved for its thermal marine environment.<sup id="cite_ref-Zillig_3-20" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Genomics">Genomics</h3></div>
<p>Construction of a physical map of <i>T. celer</i> Vu13 by way of <a href="Restriction_enzyme" title="Restriction enzyme">restriction enzyme</a> fragments revealed a length of 1,890 + 27 kilobases (kb).<sup id="cite_ref-Noll_2-2" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The molecular ratio of <a href="Guanine" title="Guanine">guanine</a> to <a href="Cytosine" title="Cytosine">cytosine</a> bases is roughly 56.6%.<sup id="cite_ref-Zillig_3-21" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> This value was determined by averaging both the <a href="GC-content" title="GC-content">GC-content</a> acquired by <a href="High-performance_liquid_chromatography" title="High-performance liquid chromatography">high-performance liquid chromatography</a> and <a href="Melting_point" title="Melting point">melting point</a> (T<sub>M</sub>) calculations.<sup id="cite_ref-Zillig_3-22" class="reference"><a href="#cite_note-Zillig-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> <i>T. celer</i> is considered to be one of the slowest evolving archaeon species, indicating that the genome could be used as a <a href="Model_organism" title="Model organism">model organism</a> for those studying early genome characteristics.<sup id="cite_ref-Noll_2-3" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p><p>In 1989, <i>T. celer</i> was the first archaeon discovered to house a circularized genome.<sup id="cite_ref-Noll_2-4" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Genome shape was determined through three separate experiments, all using <a href="Restriction_enzymes" class="mw-redirect" title="Restriction enzymes">restriction enzymes</a>.<sup id="cite_ref-Noll_2-5" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> The <i>T. celer</i> genome was digested with restriction enzymes <i>Nhe</i>, <i>Spe</i>, and <i>Xba</i>.<sup id="cite_ref-Noll_2-6" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Following digestion, hybridization analyses were used to determine genome shape.<sup id="cite_ref-Noll_2-7" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Probes were synthesized from cloned genes of the 16S rRNA and 23S rRNA.<sup id="cite_ref-Noll_2-8" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Both <i>Spe</i> and <i>Nhe</i> produced five fragments, all of similar shape and size.<sup id="cite_ref-Noll_2-9" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Digestion with <i>Xba</i> produced eight fragments.<sup id="cite_ref-Noll_2-10" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Using overlap patterns, the shape of the genome was determined to be circular.<sup id="cite_ref-Noll_2-11" class="reference"><a href="#cite_note-Noll-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Application">Application</h2></div>
<p>The domain Archaea is currently split into three major groups consisting of the extreme <a href="Thermophiles" class="mw-redirect" title="Thermophiles">thermophiles</a>, the extreme <a href="Halophiles" class="mw-redirect" title="Halophiles">halophiles</a>, and the extreme <a href="Thermophiles" class="mw-redirect" title="Thermophiles">thermophiles</a> that are able to reduce <a href="Sulfur" title="Sulfur">sulfur</a> (<a href="Methanogens" class="mw-redirect" title="Methanogens">methanogens</a>).<sup id="cite_ref-Achenbach_1-9" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> These three groups are not believed to have arisen independently, but instead evolved from one to another.<sup id="cite_ref-Achenbach_1-10" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p><p>The discovery of <i>T. celer</i> turned the position of the archaebacterial <a href="Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a>.<sup id="cite_ref-Achenbach_1-11" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> It was discovered to share a closer phylogenetic relationship with methanogenic archaebacteria, as opposed its <a href="Phenotypic" class="mw-redirect" title="Phenotypic">phenotypic</a> analogue, extremely thermophilic archaebacteria. This discovery was made through sequence analysis of the 16S rRNA and resulted in a rerooting of the phylogenetic tree.<sup id="cite_ref-Achenbach_1-12" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p><p>This discovery suggests that extreme thermophiles could be the earliest archaeon ancestor when considering their slow <a href="Evolution" title="Evolution">evolution</a> patterns, as well as the distribution of extreme thermophiles into both their own grouping, as well as that of the methanogens.<sup id="cite_ref-Achenbach_1-13" class="reference"><a href="#cite_note-Achenbach-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>
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<li id="cite_note-Achenbach-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Achenbach_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Achenbach_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Achenbach_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Achenbach_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Achenbach_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Achenbach_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Achenbach_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Achenbach_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Achenbach_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Achenbach_1-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-Achenbach_1-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-Achenbach_1-11"><sup><i><b>l</b></i></sup></a> <a href="#cite_ref-Achenbach_1-12"><sup><i><b>m</b></i></sup></a> <a href="#cite_ref-Achenbach_1-13"><sup><i><b>n</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">
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<li id="cite_note-Blamey-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-Blamey_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Blamey_5-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Blamey_5-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Blamey_5-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text">Blamey, J., M. Chiong, C. Lopez, and E. Smith. 1999. Optimization of the growth conditions of the extremely thermophilic microorganisms Thermococcus celer and <a href="Pyrococcus_woesei" title="Pyrococcus woesei">Pyrococcus woesei</a>. Journal of Microbiological methods. Vol: 38:1-2:169-175. Print.</span>
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</ol></div></div>
<div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2></div>
<ul><li><cite id="CITEREFLeeMakhatadzeWong2005" class="citation journal cs1">Lee, Chi-Fung; Makhatadze, George I.; Wong, Kam-Bo (2005). "Effects of Charge-to-Alanine Substitutions on the Stability of Ribosomal Protein L30e from Thermococcus celer". <i>Biochemistry</i>. <b>44</b> (51): <span class="nowrap">16817–</span>16825. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1021%2Fbi0519654">10.1021/bi0519654</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16363795">16363795</a>.</cite></li>
<li><cite id="CITEREFKimBaeKimKwon2011" class="citation journal cs1">Kim, Kee Pum; Bae, Heejin; Kim, In Hye; Kwon, Suk-Tae (2011). "Cloning, expression, and PCR application of DNA polymerase from the hyperthermophilic archaeon, Thermococcus celer". <i>Biotechnology Letters</i>. <b>33</b> (2): <span class="nowrap">339–</span>346. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs10529-010-0434-2">10.1007/s10529-010-0434-2</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/20953664">20953664</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a>&nbsp;<a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:20110756">20110756</a>.</cite></li>
<li><cite id="CITEREFReedLewisTrejoWinston2013" class="citation journal cs1">Reed, Christopher J.; Lewis, Hunter; Trejo, Eric; Winston, Vern; Evilia, Caryn (14 August 2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3787623">"Protein Adaptations in Archaeal Extremophiles"</a>. <i>Archaea</i>. <b>2013</b>: 14. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1155%2F2013%2F373275">10.1155/2013/373275</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&nbsp;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3787623">3787623</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24151449">24151449</a>.</cite></li>
<li><cite id="CITEREFWongBycroftWong2003" class="citation journal cs1">Wong, Kam-Bo; Bycroft, Mark; Wong, Kam-Bo (March 18, 2003). "Crystal structure of ribosomal protein L30e from the extreme thermophile Thermococcus celer: Thermal stability and RNA binding". <i>Biochemistry</i>. <b>42</b> (10): <span class="nowrap">2857–</span>65. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1021%2Fbi027131s">10.1021/bi027131s</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12627951">12627951</a>.</cite></li></ul>
<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>
<ul><li><a rel="nofollow" class="external text" href="http://bacdive.dsmz.de/index.php?search=16860&amp;submit=Search">Type strain of <i>Thermococcus celer</i> at Bac<i>Dive</i> - the Bacterial Diversity Metadatabase</a></li></ul>
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</style></div><div role="navigation" class="navbox" aria-labelledby="Taxon_identifiers2080" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers2080" style="font-size:114%;margin:0 4em">Taxon identifiers</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;"><i>Thermococcus celer</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><span style="white-space:nowrap;"><a href="Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q7783145" class="extiw external" title="wikidata:Q7783145">Q7783145</a></span></span></span></li>
<li><span style="white-space:nowrap;"><a href="Wikispecies" title="Wikispecies">Wikispecies</a>: <span class="uid"><span class="external"><a href="https://species.wikimedia.org/wiki/Thermococcus_celer" class="extiw external" title="wikispecies:Thermococcus celer">Thermococcus celer</a></span></span></span></li>
<li><span style="white-space:nowrap;"><a href="BacDive" title="BacDive">BacDive</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://bacdive.dsmz.de/strain/16860">16860</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="Catalogue_of_Life" title="Catalogue of Life">CoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.catalogueoflife.org/data/taxon/7BWV4">7BWV4</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://eol.org/pages/973272">973272</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.gbif.org/species/1000295">1000295</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&amp;id=10035243">10035243</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="Integrated_Taxonomic_Information_System" title="Integrated Taxonomic Information System">ITIS</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&amp;search_value=951938">951938</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="List_of_Prokaryotic_names_with_Standing_in_Nomenclature" title="List of Prokaryotic names with Standing in Nomenclature">LPSN</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://lpsn.dsmz.de/species/thermococcus-celer">thermococcus-celer</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&amp;id=2264">2264</a></span></span></li>
<li><span style="white-space:nowrap;"><a href="Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=1043520">1043520</a></span></span></li>
<li><span style="white-space:nowrap;">SeqCode Registry: <span class="uid"><a rel="nofollow" class="external text" href="https://seqco.de/i:21498">21498</a></span></span></li></ul>
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