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<h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Thermogenin</span></h1>
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<div id="mw-content-text" class="mw-body-content mw-content-ltr" lang="de" dir="ltr"><div class="mw-content-ltr mw-parser-output" lang="de" dir="ltr"><table class="wikitable hintergrundfarbe-basis infobox float-right" id="Vorlage_Infobox_Protein_2.A.29.3.2" style="font-size:90%; margin-top:0; width:350px;" summary="Infobox Protein">
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<th colspan="3" style="background:#90EE90; color:#202122;">uncoupling protein 1 (mitochondrial, proton carrier)
</th></tr>
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<th colspan="3" style="background:#90EE90; color:#202122;;">Eigenschaften des menschlichen Proteins
</th></tr>
<tr>
<td><a href="Molare_Masse" title="Molare Masse">Masse</a>/Länge <a href="Prim%C3%A4rstruktur" title="Primärstruktur">Primärstruktur</a>
</td>
<td colspan="2" style="text-align:center;">307 Aminosäuren
</td></tr>
<tr>
<td><a href="Sekund%C3%A4rstruktur" title="Sekundärstruktur">Sekundär-</a> bis <a href="Quart%C3%A4rstruktur" title="Quartärstruktur">Quartärstruktur</a>
</td>
<td colspan="2" style="text-align:center;">multipass Membranprotein
</td></tr>
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<th colspan="3" style="background:#90EE90; color:#202122;">Bezeichner
</th></tr>
<tr>
<td><a href="Human_Genome_Organisation" title="Human Genome Organisation">Gen-Namen</a>
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<td colspan="2" class="" style="text-align:center;"><i><a rel="nofollow" class="external text" href="https://www.genenames.org/tools/search/#!/all?query=12517">UCP1</a></i>, SLC25A7, UCP, Thermogenin
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<td>Externe IDs
</td>
<td colspan="2" class="">
<ul><li><a href="Online_Mendelian_Inheritance_in_Man" title="Online Mendelian Inheritance in Man">OMIM</a>: <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/omim/113730">113730</a></li>
<li><a href="UniProt" title="UniProt">UniProt</a> <a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprotkb/P25874">P25874</a></li>
<li><a href="Mouse_Genome_Informatics" title="Mouse Genome Informatics">MGI</a>: <a rel="nofollow" class="external text" href="https://www.informatics.jax.org/marker/MGI:98894">98894</a></li></ul>
</td></tr>
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<th colspan="3" style="background:#90EE90; color:#202122;">Transporter-Klassifikation
</th></tr>
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<td><a href="Transporter_Classification_Database" title="Transporter Classification Database">TCDB</a>
</td>
<td colspan="2" class="" style="text-align:center;"><a rel="nofollow" class="external text" href="https://tcdb.org/search/result.php?tc=2.A.29.3.2">2.A.29.3.2</a>
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<tr>
<td><a href="Membrantransport" title="Membrantransport">Bezeichnung</a>
</td>
<td colspan="2" style="text-align:center;"><a href="Mitochondrialer_Carrier" class="mw-redirect" title="Mitochondrialer Carrier">Mitochondrialer Carrier</a>
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<th colspan="3" style="background:#90EE90; color:#202122;">Vorkommen
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<td style="background:#C3FDB8; color:#202122;">Homologie-Familie
</td>
<td colspan="2" style="text-align:center;"><a rel="nofollow" class="external text" href="http://hogenom.univ-lyon1.fr/query_sequence?seq=P25874">mitochondrial carrier</a>
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<td style="background:#C3FDB8; color:#202122;">Übergeordnetes <a href="Taxon" title="Taxon">Taxon</a>
</td>
<td colspan="2" style="text-align:center;"><a href="Eukaryoten" title="Eukaryoten">Eukaryoten</a>
</td></tr>
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<td colspan="3" style="background:#90EE90; color:#202122; text-align:center;"><a href="Homologie_(Genetik)#Homologie_zwischen_verdoppelten_oder_fremden_Genen" title="Homologie (Genetik)">Orthologe</a>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;">
</td>
<td style="background:#C3FDB8; color:#202122; text-align:center;">Mensch
</td>
<td style="background:#C3FDB8; color:#202122; text-align:center;">Maus
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="Entrez_Gene" title="Entrez Gene">Entrez</a>
</td>
<td><span class=""><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=gene&cmd=retrieve&dopt=default&list_uids=7350&rn=1">7350</a></span>
</td>
<td><span class=""><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=gene&cmd=retrieve&dopt=default&list_uids=22227&rn=1">22227</a></span>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="Ensembl" title="Ensembl">Ensembl</a>
</td>
<td><span class=""><small><small><a rel="nofollow" class="external text" href="http://www.ensembl.org/Homo_sapiens/geneview?gene=ENSG00000109424;db=core">ENSG00000109424</a></small></small></span>
</td>
<td><span class=""><small><small><a rel="nofollow" class="external text" href="http://www.ensembl.org/Mus_musculus/geneview?gene=ENSMUSG00000031710;db=core">ENSMUSG00000031710</a></small></small></span>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="UniProt" title="UniProt">UniProt</a>
</td>
<td><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprotkb/P25874">P25874</a>
</td>
<td><a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprotkb/P12242">P12242</a>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">Refseq</a> (mRNA)
</td>
<td><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_021833">NM_021833</a>
</td>
<td><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NM_009463">NM_009463</a>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">Refseq</a> (Protein)
</td>
<td><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_068605">NP_068605</a>
</td>
<td><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?val=NP_033489">NP_033489</a>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="Genlocus" title="Genlocus">Genlocus</a>
</td>
<td><span class=""> <a rel="nofollow" class="external text" href="https://genome.ucsc.edu/cgi-bin/hgTracks?org=Human&position=chr4:141700500-141709457">Chr 4: 141.7 – 141.71 Mb</a> </span>
</td>
<td><span class=""> <a rel="nofollow" class="external text" href="https://genome.ucsc.edu/cgi-bin/hgTracks&position=chr8:86180457-86188557">Chr 8: 86.18 – 86.19 Mb</a> </span>
</td></tr>
<tr>
<td style="background:#C3FDB8; color:#202122;"><a href="PubMed" title="PubMed">PubMed</a>-Suche
</td>
<td><span class=""><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/sites/entrez?db=gene&cmd=Link&LinkName=gene_pubmed&from_uid=7350">7350</a></span>
</td>
<td><span class=""><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/sites/entrez?db=gene&cmd=Link&LinkName=gene_pubmed&from_uid=22227">22227</a></span>
<p><span class="editoronly" style="display:none;"></span>
</p>
</td></tr></tbody></table><p><span class="editoronly" style="display:none;"></span>
</p><p><b>Thermogenin</b> (auch <i>entkoppelndes Protein</i>, engl. <i>Uncoupling Protein 1</i> bzw. <b>UCP1</b> genannt<sup id="cite_ref-entrez_1-0" class="reference"><a href="#cite_note-entrez-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>) ist ein <a href="Transmembranprotein" title="Transmembranprotein">Transmembranprotein</a>, das sich in der <a href="Mitochondrion" class="mw-redirect" title="Mitochondrion">Mitochondrienmembran</a> des <a href="Braunes_Fettgewebe" title="Braunes Fettgewebe">braunen Fettgewebes</a> findet (bei Ratten kommt es auch im <a href="Fettgewebe#Weißes_Fettgewebe" title="Fettgewebe">weißen Fettgewebe</a> vor<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>). Durch seine Funktion kann Wärme ohne Muskelaktivität wie etwa Zittern generiert werden. Die durch UCP1 bewirkte <i><a href="Thermogenese" title="Thermogenese">Thermogenese</a></i> ist die vorherrschende Art der Wärmeerzeugung bei <a href="Winterschlaf" title="Winterschlaf">Winterschlaf</a> haltenden oder neugeborenen Säugetieren einschließlich menschlichen Säuglingen und Kleinkindern. Thermogenin gehört zu den <i><a href="Uncoupling_protein" class="mw-redirect" title="Uncoupling protein">uncoupling proteins</a></i>.
</p>
<div class="mw-heading mw-heading2"><h2 id="Wirkmechanismus">Wirkmechanismus</h2></div>
<p>Der molekulare Mechanismus der durch Thermogenin vermittelten Entkopplung ist recht gut verstanden – die Funktion dieses <a href="Ionenkanal" title="Ionenkanal">Ionenkanals</a> stellt innerhalb der <a href="Atmungskette" title="Atmungskette">Oxidativen Phosphorylierung</a> einen zusätzlichen Pfad für den Wiedereintritt von <a href="Proton_(Chemie)" title="Proton (Chemie)">Protonen</a> in die mitochondriale Matrix zur Verfügung. Wird Thermogenin aktiviert, schließt es – ähnlich wie <a href="2%2C4-Dinitrophenol#Wirkungsmechanismus" title="2,4-Dinitrophenol">2,4-Dinitrophenol</a> – den Protonen-Kreislauf kurz, der die <a href="Atmungskette" title="Atmungskette">Atmungskette</a> an den <a href="ATP-Synthase" title="ATP-Synthase">ATP-Synthase</a>-Komplex koppelt, welcher das <a href="Adenosintriphosphat" title="Adenosintriphosphat">Adenosintriphosphat</a> (ATP) für den Energiehaushalt der Zelle liefert: Durch Thermogenin wird diese Kopplung aufgehoben und die durch <a href="Zellatmung" title="Zellatmung">Zellatmung</a> gewonnene Energie ohne Bildung des Energiespeichers ATP unmittelbar in Wärme umgesetzt (<a href="Dissipation" title="Dissipation">Dissipation</a>).
</p><p>Thermogenin steht in Beziehung zu anderen mitochondrialen Stoffwechseltransportern wie dem Adenin-Nukleotid-Transporter, einem Protonenkanal in der <a href="Mitochondrium" title="Mitochondrium">inneren Mitochondrienmembran</a>, der den Transport von Protonen aus dem Membranzwischenraum in die mitochondriale Matrix vermittelt.
</p><p>UCP1 kommt ausschließlich in braunem Fettgewebe vor. Durch seine Anwesenheit erlangt dieses Gewebe seine enorme Fähigkeit zur Wärmeerzeugung.
</p><p>UCP1 wird durch <a href="Fetts%C3%A4ure" class="mw-redirect" title="Fettsäure">Fettsäuren</a> auf folgendem Weg aktiviert: Im <a href="Sympathisches_Nervensystem" class="mw-redirect" title="Sympathisches Nervensystem">sympathischen Nervensystem</a> wird <a href="Noradrenalin" title="Noradrenalin">Noradrenalin</a> freigesetzt, das an den <a href="Beta-Adrenozeptor" class="mw-redirect" title="Beta-Adrenozeptor">β3-Adrenozeptor</a> in der <a href="Zellmembran" title="Zellmembran">Zellmembran</a> der braunen <a href="Adipozyt" title="Adipozyt">Fettzellen</a> bindet. Dadurch wird eine <a href="Adenylylcyclase" class="mw-redirect" title="Adenylylcyclase">Adenylylcyclase</a> aktiviert, die die Umwandlung von ATP zu <a href="Cyclisches_Adenosinmonophosphat" title="Cyclisches Adenosinmonophosphat">cyclischem Adenosinmonophosphat</a> (cAMP) katalysiert. cAMP seinerseits aktiviert die <a href="Proteinkinase_A" title="Proteinkinase A">Proteinkinase A</a>, deren aktive C-Untereinheiten sich von den regulatorischen R-Untereinheiten lösen. Die aktive Proteinkinase A ihrerseits aktiviert durch <a href="Phosphorylierung" title="Phosphorylierung">Phosphorylierung</a> eine <a href="Lipasen" title="Lipasen">Lipase</a>. Diese Lipase katalysiert die Umwandlung von <a href="Triglyceride" title="Triglyceride">Triglyceriden</a> in freie Fettsäuren, die schließlich UCP1 aktivieren, indem sie dessen Hemmung durch <a href="Purinnukleotide" class="mw-redirect" title="Purinnukleotide">Purinnukleotide</a> (<a href="Guanosindiphosphat" title="Guanosindiphosphat">GDP</a>, <a href="Adenosindiphosphat" title="Adenosindiphosphat">ADP</a>) aufheben. Die Thermogenese wird dadurch beendet, dass die Mitochondrien freie Fettsäurereste oxidieren. Dadurch wird UCP1 inaktiv und die Zelle kehrt zur normalen Zellatmung zurück, bei der ATP synthetisiert wird.
</p>
<div class="mw-heading mw-heading2"><h2 id="Entdeckungsgeschichte">Entdeckungsgeschichte</h2></div>
<p>Entdeckt wurde der Ionenkanal 1979 als <i>Uncoupling Protein</i><sup id="cite_ref-pmid348493_3-0" class="reference"><a href="#cite_note-pmid348493-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>, 1988 wurde es zum ersten Mal kloniert.<sup id="cite_ref-pmid3410843_4-0" class="reference"><a href="#cite_note-pmid3410843-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid3202878_5-0" class="reference"><a href="#cite_note-pmid3202878-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>
</p><p>Ein zu Thermogenin homologes <a href="Gen" title="Gen">Gen</a> konnte 1997 identifiziert werden und wurde UCP2 genannt. In den letzten zehn Jahren wurden drei weitere UCP1-Homologe gefunden, darunter UCP3, UCP4, und BMCP1 (auch als UCP5 bekannt). Erst in den vergangenen Jahren der Forschung konnte braunes Fettgewebe auch bei erwachsenen Menschen gefunden werden.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Siehe_auch">Siehe auch</h2></div>
<ul><li><a href="Leptin" title="Leptin">Leptin</a></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Literatur">Literatur</h2></div>
<ul><li>David Nelson, Michael Cox: <i>Lehninger Principles of Biochemistry</i>. 4. Edition. W.H. Freeman, New York 2005, ISBN 0-7167-4339-6</li>
<li>Ricquier D, Bouillaud F: <cite class="lang" lang="en" dir="auto" style="font-style:italic">The uncoupling protein homologues: UCP1, UCP2, UCP3, StUCP and AtUCP</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic"><a href="Biochem._J." class="mw-redirect" title="Biochem. J.">Biochem. J.</a></cite> 345 Pt 2. Jahrgang, 2000, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>161–79</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/10620491?dopt=Abstract">PMID 10620491</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=The+uncoupling+protein+homologues%3A+UCP1%2C+UCP2%2C+UCP3%2C+StUCP+and+AtUCP&rft.au=Ricquier+D%2C+Bouillaud+F&rft.btitle=Biochem.+J.&rft.date=2000&rft.genre=book&rft.pages=161-79&rft.pmid=10620491&rft.volume=345+Pt+2.+Jahrgang" style="display:none"> </span></li>
<li>Muzzin P: <cite class="lang" lang="en" dir="auto" style="font-style:italic">The uncoupling proteins</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Ann. Endocrinol. (Paris)</cite>. 63. Jahrgang, 2 Pt 1, 2002, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>106–10</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/11994670?dopt=Abstract">PMID 11994670</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=The+uncoupling+proteins&rft.au=Muzzin+P&rft.date=2002&rft.genre=journal&rft.issue=2+Pt+1&rft.jtitle=Ann.+Endocrinol.+%28Paris%29&rft.pages=106-10&rft.pmid=11994670&rft.volume=63.+Jahrgang" style="display:none"> </span></li>
<li>Del Mar Gonzalez-Barroso M, Ricquier D, Cassard-Doulcier AM: <cite class="lang" lang="en" dir="auto" style="font-style:italic">The human uncoupling protein-1 gene (UCP1): present status and perspectives in obesity research</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic"><a href="Obesity_Reviews" title="Obesity Reviews">Obesity Reviews</a></cite>. 1. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>2</span>, 2002, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>61–72</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/12119988?dopt=Abstract">PMID 12119988</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=The+human+uncoupling+protein-1+gene+%28UCP1%29%3A+present+status+and+perspectives+in+obesity+research&rft.au=Del+Mar+Gonzalez-Barroso+M%2C+Ricquier+D%2C+Cassard-Doulcier+AM&rft.date=2002&rft.genre=journal&rft.issue=2&rft.jtitle=Obesity+Reviews&rft.pages=61-72&rft.pmid=12119988&rft.volume=1.+Jahrgang" style="display:none"> </span></li>
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<li>Rim JS, Kozak LP: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Regulatory motifs for CREB-binding protein and Nfe2l2 transcription factors in the upstream enhancer of the mitochondrial uncoupling protein 1 gene</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic">J. Biol. Chem.</cite> 277. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>37</span>, 2002, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>34589–600</span>, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1074/jbc.M108866200">10.1074/jbc.M108866200</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/12084707?dopt=Abstract">PMID 12084707</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=Regulatory+motifs+for+CREB-binding+protein+and+Nfe2l2+transcription+factors+in+the+upstream+enhancer+of+the+mitochondrial+uncoupling+protein+1+gene&rft.au=Rim+JS%2C+Kozak+LP&rft.date=2002&rft.doi=10.1074%2Fjbc.M108866200&rft.genre=journal&rft.issue=37&rft.jtitle=J.+Biol.+Chem.&rft.pages=34589-600&rft.pmid=12084707&rft.volume=277.+Jahrgang" style="display:none"> </span></li>
<li>Kieć-Wilk B, Wybrańska I, Malczewska-Malec M, <i>et al.</i>: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Correlation of the -3826A >G polymorphism in the promoter of the uncoupling protein 1 gene with obesity and metabolic disorders in obese families from southern Poland</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic">J. Physiol. Pharmacol.</cite> 53. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>3</span>, 2003, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>477–90</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/12375583?dopt=Abstract">PMID 12375583</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=Correlation+of+the+-3826A+%3EG+polymorphism+in+the+promoter+of+the+uncoupling+protein+1+gene+with+obesity+and+metabolic+disorders+in+obese+families+from+southern+Poland&rft.au=Kie%C4%87-Wilk+B%2C+Wybra%C5%84ska+I%2C+Malczewska-Malec+M%2C+...&rft.date=2003&rft.genre=journal&rft.issue=3&rft.jtitle=J.+Physiol.+Pharmacol.&rft.pages=477-90&rft.pmid=12375583&rft.volume=53.+Jahrgang" style="display:none"> </span></li>
<li>Strausberg RL, Feingold EA, Grouse LH, <i>et al.</i>: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Generation and initial analysis of more than 15,000 full-length human and mouse cDNA sequences</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic"><a href="Proc._Natl._Acad._Sci._USA" class="mw-redirect" title="Proc. Natl. Acad. Sci. USA">Proc. Natl. Acad. Sci. USA</a></cite>. 99. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>26</span>, 2003, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>16899–903</span>, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1073/pnas.242603899">10.1073/pnas.242603899</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/12477932?dopt=Abstract">PMID 12477932</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=Generation+and+initial+analysis+of+more+than+15%2C000+full-length+human+and+mouse+cDNA+sequences&rft.au=Strausberg+RL%2C+Feingold+EA%2C+Grouse+LH%2C+...&rft.date=2003&rft.doi=10.1073%2Fpnas.242603899&rft.genre=journal&rft.issue=26&rft.jtitle=Proc.+Natl.+Acad.+Sci.+USA&rft.pages=16899-903&rft.pmid=12477932&rft.volume=99.+Jahrgang" style="display:none"> </span></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Weblinks">Weblinks</h2></div>
<ul><li><a rel="nofollow" class="external text" href="http://news.bbc.co.uk/2/hi/health/5335176.stm">Seaweed anti-obesity tablet hope (BBC-News - Zusammenspiel von Flucoxanthin und UCP1, engl.)</a></li>
<li><a rel="nofollow" class="external text" href="http://www.nlm.nih.gov/cgi/mesh/2007/MB_cgi?mode=&index=54059&field=all&HM=&II=&PA=&form=&input=">MeshName</a></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Einzelnachweise">Einzelnachweise</h2></div>
<ol class="references">
<li id="cite_note-entrez-1"><span class="mw-cite-backlink"><a href="#cite_ref-entrez_1-0">↑</a></span> <span class="reference-text"><span class="cite"><a rel="nofollow" class="external text" href="http://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=7350"><i>Entrez Gene: UCP1 uncoupling protein 1 (mitochondrial, proton carrier).</i></a><span class="Abrufdatum"> Abgerufen am 25. Februar 2011</span> (englisch).</span><span style="display: none;" class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Adc&rfr_id=info%3Asid%2Fde.wikipedia.org%3AThermogenin&rft.title=Entrez+Gene%3A+UCP1+uncoupling+protein+1+%28mitochondrial%2C+proton+carrier%29&rft.description=Entrez+Gene%3A+UCP1+uncoupling+protein+1+%28mitochondrial%2C+proton+carrier%29&rft.identifier=&rft.date=&rft.language=en"> </span></span>
</li>
<li id="cite_note-2"><span class="mw-cite-backlink"><a href="#cite_ref-2">↑</a></span> <span class="reference-text">Wang LH, e.a.: <i>Electroacupuncture Relieves Obesity by Up-regulating PGC-1 α/UCP-1 Signaling in White Adipose Tissue in Diet-induced Obesity Rats</i>. Zhen Ci Yan Jiu. 2018 Aug 25;43(8):495-500. Chinese. <a href="https://doi.org/10.13702/j.1000-0607.170744" class="extiw external" title="doi:10.13702/j.1000-0607.170744">doi:10.13702/j.1000-0607.170744</a>. <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/30232852?dopt=Abstract">PMID 30232852</a>.</span>
</li>
<li id="cite_note-pmid348493-3"><span class="mw-cite-backlink"><a href="#cite_ref-pmid348493_3-0">↑</a></span> <span class="reference-text">Nicholls DG, Bernson VS, Heaton GM: <cite class="lang" lang="en" dir="auto" style="font-style:italic">The identification of the component in the inner membrane of brown adipose tissue mitochondria responsible for regulating energy dissipation</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Experientia Suppl.</cite> 32. Jahrgang, 1978, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>89–93</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/348493?dopt=Abstract">PMID 348493</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=The+identification+of+the+component+in+the+inner+membrane+of+brown+adipose+tissue+mitochondria+responsible+for+regulating+energy+dissipation&rft.au=Nicholls+DG%2C+Bernson+VS%2C+Heaton+GM&rft.btitle=Experientia+Suppl.&rft.date=1978&rft.genre=book&rft.pages=89-93&rft.pmid=348493&rft.volume=32.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid3410843-4"><span class="mw-cite-backlink"><a href="#cite_ref-pmid3410843_4-0">↑</a></span> <span class="reference-text">Kozak LP, Britton JH, Kozak UC, Wells JM: <cite class="lang" lang="en" dir="auto" style="font-style:italic">The mitochondrial uncoupling protein gene. Correlation of exon structure to transmembrane domains</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic">J. Biol. Chem.</cite> 263. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>25</span>, 1988, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>12274–7</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/3410843?dopt=Abstract">PMID 3410843</a> (englisch, <a rel="nofollow" class="external text" href="https://www.jbc.org/article/S0021-9258(18)37751-2/pdf">jbc.org</a> [PDF; abgerufen am 20. Februar 2008]).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=The+mitochondrial+uncoupling+protein+gene.+Correlation+of+exon+structure+to+transmembrane+domains&rft.au=Kozak+LP%2C+Britton+JH%2C+Kozak+UC%2C+...&rft.date=1988&rft.genre=journal&rft.issue=25&rft.jtitle=J.+Biol.+Chem.&rft.pages=12274-7&rft.pmid=3410843&rft.volume=263.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid3202878-5"><span class="mw-cite-backlink"><a href="#cite_ref-pmid3202878_5-0">↑</a></span> <span class="reference-text">Bouillaud F, Raimbault S, Ricquier D: <cite class="lang" lang="en" dir="auto" style="font-style:italic">The gene for rat uncoupling protein: complete sequence, structure of primary transcript and evolutionary relationship between exons</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic"><a href="Biochem._Biophys._Res._Commun." class="mw-redirect" title="Biochem. Biophys. Res. Commun.">Biochem. Biophys. Res. Commun.</a></cite> 157. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>2</span>, 1988, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>783–92</span>, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1016/S0006-291X%2888%2980318-8">10.1016/S0006-291X(88)80318-8</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/3202878?dopt=Abstract">PMID 3202878</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=The+gene+for+rat+uncoupling+protein%3A+complete+sequence%2C+structure+of+primary+transcript+and+evolutionary+relationship+between+exons&rft.au=Bouillaud+F%2C+Raimbault+S%2C+Ricquier+D&rft.date=1988&rft.doi=10.1016%2FS0006-291X%2888%2980318-8&rft.genre=journal&rft.issue=2&rft.jtitle=Biochem.+Biophys.+Res.+Commun.&rft.pages=783-92&rft.pmid=3202878&rft.volume=157.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-6"><span class="mw-cite-backlink"><a href="#cite_ref-6">↑</a></span> <span class="reference-text">Jan Nedergaard, Tore Bengtsson, Barbara Cannon: <cite style="font-style:italic">Unexpected evidence for active brown adipose tissue in adult humans</cite>. In: <cite style="font-style:italic">American Journal of Physiology. Endocrinology and Metabolism</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em"> </span>293</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>2</span>, August 2007, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a> <span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%220193-1849%22&key=cql">0193-1849</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>E444–452</span>, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.1152/ajpendo.00691.2006">10.1152/ajpendo.00691.2006</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/17473055?dopt=Abstract">PMID 17473055</a>.<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=Unexpected+evidence+for+active+brown+adipose+tissue+in+adult+humans&rft.au=Jan+Nedergaard%2C+Tore+Bengtsson%2C+Barbara+Cannon&rft.date=2007-08&rft.doi=10.1152%2Fajpendo.00691.2006&rft.genre=journal&rft.issn=0193-1849&rft.issue=2&rft.jtitle=American+Journal+of+Physiology.+Endocrinology+and+Metabolism&rft.pages=E444-452&rft.pmid=17473055&rft.volume=293" style="display:none"> </span></span>
</li>
<li id="cite_note-7"><span class="mw-cite-backlink"><a href="#cite_ref-7">↑</a></span> <span class="reference-text">Masayuki Saito, Yuko Okamatsu-Ogura, Mami Matsushita, Kumiko Watanabe, Takeshi Yoneshiro: <cite style="font-style:italic">High incidence of metabolically active brown adipose tissue in healthy adult humans: effects of cold exposure and adiposity</cite>. In: <cite style="font-style:italic">Diabetes</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em"> </span>58</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>7</span>, Juli 2009, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a> <span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%221939-327X%22&key=cql">1939-327X</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>1526–1531</span>, <a href="Digital_Object_Identifier" title="Digital Object Identifier">doi</a>:<span class="uri-handle" style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://doi.org/10.2337/db09-0530">10.2337/db09-0530</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/19401428?dopt=Abstract">PMID 19401428</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2699872/">PMC 2699872</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rfr_id=info:sid/de.wikipedia.org:Thermogenin&rft.atitle=High+incidence+of+metabolically+active+brown+adipose+tissue+in+healthy+adult+humans%3A+effects+of+cold+exposure+and+adiposity&rft.au=Masayuki+Saito%2C+Yuko+Okamatsu-Ogura%2C+Mami+Matsushita%2C+...&rft.date=2009-07&rft.doi=10.2337%2Fdb09-0530&rft.genre=journal&rft.issn=1939-327X&rft.issue=7&rft.jtitle=Diabetes&rft.pages=1526-1531&rft.pmc=2699872&rft.pmid=19401428&rft.volume=58" style="display:none"> </span></span>
</li>
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