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Thermococcus
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<h2 id="phylogeny">Phylogeny</h2> <table><tbody><tr><th colspan="1">16S rRNA based <a href="/facts/The_All-Species_Living_Tree_Project/txMadzJv">LTP</a>_08_2023<a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a><a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a><a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a></th><th colspan="1">53 marker proteins based <a href="/facts/Genome_Taxonomy_Database/hnEgBvVx">GTDB</a> 09-RS220<a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a><a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a><a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a></th></tr><tr><td><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td><i>Thermococcus</i></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. aggregans</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. aegaeus</i> Arab et al. 2000</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_alcaliphilus/BewlDHDa">T. alcaliphilus</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_litoralis/drxGRFLz">T. litoralis</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. argininiproducens</i> Park et al. 2023</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_sibiricus/s16RuPP3">T. sibiricus</a></i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td>species‑group 2</td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Pyrococcus/caLcg1wY">Pyrococcus</a></i></p></td></tr><tr><td></td></tr><tr><td><i>Thermococcus</i> s.s.</td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_barophilus/2rS4yCL5">T. barophilus</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. paralvinellae</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. acidaminovorans</i> Dirmeier et al. 2001</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. gorgonarius</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. fumicolans</i> Godfroy et al. 1996</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. pacificus</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. waiotapuensis</i> Gonzlez et al. 2001</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. zilligii</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. guaymasensis</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. eurythermalis</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. henrietii</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. nautili</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_gammatolerans/Nb2Ua4oT">T. gammatolerans</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_kodakarensis/gCVBrUQk">T. kodakarensis</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_peptonophilus/0u5Ng7xR">T. peptonophilus</a></i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_stetteri/2vkYCaGu">T. stetteri</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. cleftensis</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. siculi</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. indicus</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. celericrescens</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. aciditolerans</i> Li et al. 2021</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. camini</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_profundus/MGcN5SFD">T. profundus</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. piezophilus</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. coalescens</i> Kuwabara et al. 2005</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. prieurii</i> Gorlas et al. 2013</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. thioreducens</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_hydrothermalis/8lmMRvRg">T. hydrothermalis</a></i> Godfroy et al. 1997</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_barossii/s16RuPP3">T. barossii</a></i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. atlanticus</i> Cambon-Bonavita et al. 2004</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_celer/RjcsdmCe">T. celer</a></i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td><td><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td><i>Thermococcus</i></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. argininiproducens</i> Park et al. 2023</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. sibiricus</i> Miroshnichenko et al. 2001</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. aggregans</i> Canganella et al. 1998</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>"<i>T. bergensis</i>" Birkeland et al. 2021</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_alcaliphilus/BewlDHDa">T. alcaliphilus</a></i> Keller et al. 1997</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_litoralis/drxGRFLz">T. litoralis</a></i> Neuner et al. 2001</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td>species‑group 2</td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td><i>Thermococcus</i></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_barophilus/2rS4yCL5">T. barophilus</a></i> Marteinsson et al. 1999</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. paralvinellae</i> Hensley et al. 2014</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td>species‑group 3</td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Pyrococcus/caLcg1wY">Pyrococcus</a></i></p></td></tr><tr><td></td></tr><tr><td><i>Thermococcus</i> s.s.</td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_profundus/MGcN5SFD">T. profundus</a></i> Kobayashi and Horikoshi 1995</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. gorgonarius</i> Miroshnichenko et al. 1998</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. zilligii</i> Ronimus et al. 1999</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_stetteri/2vkYCaGu">T. stetteri</a></i> Miroshnichenko 1990</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_kodakarensis/gCVBrUQk">T. kodakarensis</a></i> Atomi et al. 2005</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_peptonophilus/0u5Ng7xR">T. peptonophilus</a></i> González et al. 1996</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_gammatolerans/Nb2Ua4oT">T. gammatolerans</a></i> Jolivet et al. 2003</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. guaymasensis</i> Canganella et al. 1998</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. eurythermalis</i> Zhao et al. 2015</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. henrietii</i> Alain et al. 2021</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. nautili</i> Soler et al. 2007</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. cleftensis</i> Hensley et al. 2014</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. pacificus</i> Miroshnichenko et al. 1998</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. siculi</i> Grote et al. 2000</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. indicus</i> Lim et al. 2021</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. camini</i> Courtine et al. 2021</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. celericrescens</i> Kuwabara et al. 2007</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>"<i>T. onnurineus</i>" Bae et al. 2006</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. piezophilus</i> Dalmasso et al. 2017</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i><a href="/facts/Thermococcus_celer/RjcsdmCe">T. celer</a></i> Zillig 1983 (type sp.)</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><i>T. barossii</i> Duffaud et al. 2005</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>"<i>T. radiotolerans</i>" Jolivet et al. 2004</p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><i>T. thioreducens</i> Pikuta et al. 2007</p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr></tbody></table> <p>Unassigned species: </p> <ul><li><i>T. coalescens</i> Kuwabara et al. 2005</li> <li><i>T. marinus</i> Jolivet et al. 2004</li> <li><i>T. mexicalis</i> Antoine 1996</li> <li><i>T. thermotolerans</i> Yang et al. 2023</li> <li>"<i>T. waimanguensis</i>" Goetz & Morgan 1999</li></ul> <h2 id="metabolism">Metabolism</h2> <p>Metabolically, <i>Thermococcus</i> spp. have developed a different form of glycolysis from eukaryotes and prokaryotes.<a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a><a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a> One example of a metabolic pathway for these organisms is the metabolism of peptides,<a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a> which occurs in three steps: first, hydrolysis of the peptides to amino acids is catalyzed by peptidases,<a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a> then the conversion of the amino acids to keto acids is catalyzed by aminotransferases,<a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a> and finally CO2 is released from the oxidative decarboxylation or the keto acids by four different enzymes,<a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a> which produces coenzyme A derivatives that are used in other important metabolic pathways.<a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a> <i>Thermococcus</i> species also have the enzyme rubisco (ribulose-1,5-bisphosphate carboxylase/oxygenase),<a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a> which is made from enzymes involved in the metabolism of nucleic acids in <i>Thermococcus kodakarensis</i>,<a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a><a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a><a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a> showing how integrated these metabolic systems truly are for these hyperthermophilic microorganisms.<a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a> Some nutrients are limiting in <i>Thermococcus</i> cell growth.<a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> Nutrients that affect cell growth the most in thermococcal species are carbon and nitrogen sources.<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a> Since thermococcal species do not metabolically generate all necessary amino acids, some have to be provided by the environment in which these organisms thrive. Some of these needed amino acids are leucine, isoleucine, and valine (the branched-chain amino acids).<a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a> When <i>Thermococcus</i> species are supplemented with these amino acids, they can metabolize them and produce acetyl-CoA or succinyl-CoA,<a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a> which are important precursors used in other metabolic pathways essential for cellular growth and respiration.<a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a> <i>Thermococcus onnurineus</i> lacks the <a href="/facts/Gene/e1swwPY6">genes</a> for <a href="/facts/Purine/9ogLp0Wc">purine</a> <a href="/facts/Nucleotide/pQKEwlJI">nucleotide</a> biosynthesis and thus relies on environmental sources to meet its purine requirements.<a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> With today's technology, <i>Thermococcus</i> members are relatively easy to grow in labs,<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a> and are therefore considered model organisms for studying the physiological and molecular pathways of extremophiles.<a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a><a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> <i>Thermococcus kodakarensis</i> is one example of a model <i>Thermococcus</i> species, a microorganism in which has had its entire genome examined and replicated.<a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a><a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a><a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> </p> <h2 id="ecology">Ecology</h2> <p>Thermococcal species can grow between 60 and 102 °C, optimal temperature at 85 °C which gives them a great ecological advantage to be the first organisms to colonize new hydrothermal environments.<a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a><a class="footnote-ref" id="fnref:58" href="#fn:58"><sup>58</sup></a><a class="footnote-ref" id="fnref:59" href="#fn:59"><sup>59</sup></a> As hyperthermophiles, there is a need for extreme environmental conditions, including temperature, pH, and salt. These conditions lead to the production of stress proteins and molecular chaperones that protect DNA as well as housekeeping cellular machinery. Thermococcus also thrives under gluconeogenic conditions. Some thermococcal species produce CO2, H2, and H2S as products of metabolism and respiration.<a class="footnote-ref" id="fnref:60" href="#fn:60"><sup>60</sup></a> The releases of these molecules are then used by other autotrophic species, aiding the diversity of hydrothermal microbial communities.<a class="footnote-ref" id="fnref:61" href="#fn:61"><sup>61</sup></a> This type of continuous enrichment culture plays a crucial role in the ecology of deep-sea hydrothermal vents,<a class="footnote-ref" id="fnref:62" href="#fn:62"><sup>62</sup></a> suggesting that thermococci interact with other organisms via metabolite exchange, which supports the growth of autotrophs.<a class="footnote-ref" id="fnref:63" href="#fn:63"><sup>63</sup></a> <i>Thermococcus</i> species that release H2 with the use of multiple hydrogenases (including CO-dependent hydrogenases) have been regarded as potential biocatalysts for water-gas shift reactions.<a class="footnote-ref" id="fnref:64" href="#fn:64"><sup>64</sup></a> </p> <h2 id="transportation-mechanisms">Transportation mechanisms</h2> <p><i>Thermococcus</i> species are naturally competent in taking up DNA and incorporating donor DNA into their genomes via homologous recombination.<a class="footnote-ref" id="fnref:65" href="#fn:65"><sup>65</sup></a> These species can produce membrane vesicles (MVs),<a class="footnote-ref" id="fnref:66" href="#fn:66"><sup>66</sup></a> formed by budding from the outermost cellular membranes,<a class="footnote-ref" id="fnref:67" href="#fn:67"><sup>67</sup></a><a class="footnote-ref" id="fnref:68" href="#fn:68"><sup>68</sup></a> which can capture and obtain plasmids from neighboring Archaea species to transfer the DNA into either themselves or surrounding species.<a class="footnote-ref" id="fnref:69" href="#fn:69"><sup>69</sup></a> These MVs are secreted from the cells in clusters, forming nanospheres or nanotubes,<a class="footnote-ref" id="fnref:70" href="#fn:70"><sup>70</sup></a> keeping the internal membranes continuous.<a class="footnote-ref" id="fnref:71" href="#fn:71"><sup>71</sup></a> Competence for DNA transfer and integration of donor DNA into the recipient genome by <a href="/facts/Homologous_recombination/WU0IIVVv">homologous recombination</a> is common in the <a href="/facts/Archaea/Adxh0aHw">archaea</a> and appears to be an adaptation for <a href="/facts/DNA_repair/hsURuYYq">repairing DNA damage</a> in the recipient cells (see <a href="/facts/Archaea/Adxh0aHw">Archaea</a> subsection "Gene transfer and genetic exchange"). </p><p><i>Thermococcus</i> species produce numerous MVs, transferring DNA, metabolites, and even toxins in some species;<a class="footnote-ref" id="fnref:72" href="#fn:72"><sup>72</sup></a> moreover, these MVs protect their contents against thermodegradation by transferring these macromolecules in a protected environment.<a class="footnote-ref" id="fnref:73" href="#fn:73"><sup>73</sup></a><a class="footnote-ref" id="fnref:74" href="#fn:74"><sup>74</sup></a> MVs also prevent infections by capturing viral particles.<a class="footnote-ref" id="fnref:75" href="#fn:75"><sup>75</sup></a> Along with transporting macromolecules, <i>Thermococcus</i> species use MVs to communicate to each other.<a class="footnote-ref" id="fnref:76" href="#fn:76"><sup>76</sup></a> Furthermore, these MVs are used by a specific species (<i>Thermococcus coalescens</i>) to indicate when aggregation should occur,<a class="footnote-ref" id="fnref:77" href="#fn:77"><sup>77</sup></a> so these typically single-celled miroorganisms can fuse into one massive single cell.<a class="footnote-ref" id="fnref:78" href="#fn:78"><sup>78</sup></a> </p><p>It has been reported that <i>Thermococcus kodakarensis</i> has four virus-like integrated gene elements containing subtilisin-like serine protease precursors.<a class="footnote-ref" id="fnref:79" href="#fn:79"><sup>79</sup></a> To date, only two viruses have been isolated from <i>Thermococcus</i> spp., PAVE1 and TPV1.<a class="footnote-ref" id="fnref:80" href="#fn:80"><sup>80</sup></a> These viruses exist in their hosts in a carrier state.<a class="footnote-ref" id="fnref:81" href="#fn:81"><sup>81</sup></a> The process of DNA replication and elongation has been extensively studied in <i>T. kodakarensis</i>.<a class="footnote-ref" id="fnref:82" href="#fn:82"><sup>82</sup></a> The DNA molecule is a circular structure consisting of about 2 million base pairs in length, and has more than 2,000 sequences that code for proteins.<a class="footnote-ref" id="fnref:83" href="#fn:83"><sup>83</sup></a> </p> <h2 id="future-technology">Future technology</h2> <p>An enzyme from <i>Thermococcus</i>, Tpa-S DNA polymerase, has been found to be more efficient in long and rapid <a href="/facts/Polymerase_chain_reaction/G0jWzJy1">polymerase chain reaction</a> (PCR) than <a href="/facts/Taq_polymerase/ElSRbPss">Taq polymerase</a>.<a class="footnote-ref" id="fnref:84" href="#fn:84"><sup>84</sup></a> Tk-SP, another enzyme from <i>T. kodakarensis</i>,<a class="footnote-ref" id="fnref:85" href="#fn:85"><sup>85</sup></a><a class="footnote-ref" id="fnref:86" href="#fn:86"><sup>86</sup></a> can degrade abnormal <a href="/facts/Prion/UJfG9EfS">prion</a> proteins (PrPSc);<a class="footnote-ref" id="fnref:87" href="#fn:87"><sup>87</sup></a> prions are misfolded proteins that can cause fatal diseases in all organisms.<a class="footnote-ref" id="fnref:88" href="#fn:88"><sup>88</sup></a> Tk-SP shows broad substrate specificity, and degraded prions exponentially in the lab setting.<a class="footnote-ref" id="fnref:89" href="#fn:89"><sup>89</sup></a> This enzyme does not require <a href="/facts/Calcium/hf252CC0">calcium</a> or any other substrate to fold, so is showing great potential in studies this far.<a class="footnote-ref" id="fnref:90" href="#fn:90"><sup>90</sup></a> Additional studies have been coordinated on the <a href="/facts/Phosphoserine_phosphatase/o5WR6oJs">phosphoserine phosphatase</a> (PSP) enzyme of <i>T. onnurineus</i>, which provided an essential component in the regulation of PSP activity.<a class="footnote-ref" id="fnref:91" href="#fn:91"><sup>91</sup></a> This information is useful for drug companies, because abnormal PSP activity leads to a major decrease in <a href="/facts/Serine/4duVl30B">serine</a> levels of the <a href="/facts/Nervous_system/CDxRKU2U">nervous system</a>, causing neurological diseases and complications.<a class="footnote-ref" id="fnref:92" href="#fn:92"><sup>92</sup></a> </p><p><i>Thermococcus</i> spp. can increase <a href="/facts/Gold_mining/9Uslr31T">gold mining</a> efficiency up to 95% due to their specific abilities in <a href="/facts/Bioleaching/1FPuznKK">bioleaching</a>.<a class="footnote-ref" id="fnref:93" href="#fn:93"><sup>93</sup></a> </p> <h2 id="see-also">See also</h2> <ul><li><a href="/facts/List_of_Archaea_genera/Cmf7AgQa">List of Archaea genera</a></li></ul> <h2 id="further-reading">Further reading</h2> <ul><li>Judicial Commission of the International Committee on Systematics of Prokaryotes (January 2005). <a href="https://doi.org/10.1099%2Fijs.0.63548-0">"The nomenclatural types of the orders Acholeplasmatales, Halanaerobiales, Halobacteriales, Methanobacteriales, Methanococcales, Methanomicrobiales, Planctomycetales, Prochlorales, Sulfolobales, Thermococcales, Thermoproteales and Verrucomicrobiales are the genera Acholeplasma, Halanaerobium, Halobacterium, Methanobacterium, Methanococcus, Methanomicrobium, Planctomyces, Prochloron, Sulfolobus, Thermococcus, Thermoproteus and Verrucomicrobium, respectively. Opinion 79"</a>. <i>International Journal of Systematic and Evolutionary Microbiology</i>. 55 (Pt 1): 517–518. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1099%2Fijs.0.63548-0">10.1099/ijs.0.63548-0</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/15653928">15653928</a>.</li> <li>Mora M, Bellack A, Ugele M, Hopf J, Wirth R (August 2014). <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4148812">"The temperature gradient-forming device, an accessory unit for normal light microscopes to study the biology of hyperthermophilic microorganisms"</a>. <i>Applied and Environmental Microbiology</i>. 80 (15): 4764–70. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/2014ApEnM..80.4764M">2014ApEnM..80.4764M</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1128%2FAEM.00984-14">10.1128/AEM.00984-14</a>. <a href="/facts/PMC_(identifier)/dX1zMt71">PMC</a> <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4148812">4148812</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/24858087">24858087</a>.</li> <li>Zillig W, Holz I, Klenk HP, Trent J, Wunderl S, Janekovic D, Imsel E, Haas B (1987). "Pyrococcus woesei, sp. nov., an ultra-thermophilic marine Archaebacterium, representing a novel order, Thermococcales". <i>Syst. Appl. Microbiol</i>. 9 (1–2): 62–70. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1987SyApM...9...62Z">1987SyApM...9...62Z</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS0723-2020%2887%2980057-7">10.1016/S0723-2020(87)80057-7</a>.</li> <li>Zillig W, Holz L, Janekovic D, Schafer W, Reiter WD (1983). "The archaebacterium <i>Thermococcus celer</i> represents a novel genus within the thermophilic branch of the archaebacteria". <i>Syst. Appl. Microbiol</i>. 4 (1): 88–94. <a href="/facts/Bibcode_(identifier)/9HtdQSGB">Bibcode</a>:<a href="https://ui.adsabs.harvard.edu/abs/1983SyApM...4...88Z">1983SyApM...4...88Z</a>. <a href="/facts/Doi_(identifier)/muM9Etpq">doi</a>:<a href="https://doi.org/10.1016%2FS0723-2020%2883%2980036-8">10.1016/S0723-2020(83)80036-8</a>. <a href="/facts/PMID_(identifier)/JlHAvMHt">PMID</a> <a href="https://pubmed.ncbi.nlm.nih.gov/23196302">23196302</a>.</li></ul> <h2 id="external-links">External links</h2> <ul><li>John R. Stevenson. <a href="http://www.cas.muohio.edu/~stevenjr/mbi202/evolution202.html">"Microbial Evolution and Systematics"</a>. <i>General Microbiology II</i>. Retrieved 2008-07-13.</li> <li><a href="http://bacdive.dsmz.de/index.php?search=Thermococcus&submit=Search"><i>Thermococcus</i> at Bac<i>Dive</i> - the Bacterial Diversity Metadatabase</a></li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>See the NCBI webpage on Thermococcus. Data extracted from the "NCBI taxonomy resources". National Center for Biotechnology Information. Retrieved 2007-03-19. <a href="/wiki/National_Center_for_Biotechnology_Information" target="_blank">/wiki/National_Center_for_Biotechnology_Information</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Canganella F, Jones WJ, Gambacorta A, Antranikian G (October 1998). "Thermococcus guaymasensis sp. nov. and Thermococcus aggregans sp. nov., two novel thermophilic archaea isolated from the Guaymas Basin hydrothermal vent site". International Journal of Systematic Bacteriology. 48 Pt 4 (4): 1181–5. doi:10.1099/00207713-48-4-1181. PMID 9828419. <a href="https://doi.org/10.1099%2F00207713-48-4-1181" target="_blank">https://doi.org/10.1099%2F00207713-48-4-1181</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Tagashira K, Fukuda W, Matsubara M, Kanai T, Atomi H, Imanaka T (January 2013). 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International Journal of Systematic Bacteriology. 48 Pt 4 (4): 1181–5. doi:10.1099/00207713-48-4-1181. PMID 9828419. <a href="https://doi.org/10.1099%2F00207713-48-4-1181" target="_blank">https://doi.org/10.1099%2F00207713-48-4-1181</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>Amenábar MJ, Flores PA, Pugin B, Boehmwald FA, Blamey JM (2013). "Archaeal diversity from hydrothermal systems of Deception Island, Antarctica". Polar Biology. 36 (3): 373–380. Bibcode:2013PoBio..36..373A. doi:10.1007/s00300-012-1267-3. S2CID 11705986. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Krupovic M, Gonnet M, Hania WB, Forterre P, Erauso G (2013). "Insights into dynamics of mobile genetic elements in hyperthermophilic environments from five new Thermococcus plasmids". PLOS ONE. 8 (1): e49044. 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