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data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> </div> <div id="siteSub" class="noprint">Da Wikipedia, l'enciclopedia libera.</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="it" dir="ltr"><table class="infobox sinottico" summary="Tabella sinottica che riassume i principali dati del soggetto" style="width:230px"><tbody><tr class="sinottico_testata"><th colspan="2" style="background-color: lightblue; color: black;"><div class="noprint" style="position:relative; float:right; font-size:70%;"><figure class="mw-default-size noresize" typeof="mw:File"><span title="Come leggere il tassobox"><img alt="Come leggere il tassobox" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Information-silk.svg/16px-Information-silk.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/51/Information-silk.svg/24px-Information-silk.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/51/Information-silk.svg/32px-Information-silk.svg.png 2x" data-file-width="16" data-file-height="16" usemap="#ImageMap_37fb70b5d7ceee19" /></span><map name="ImageMap_37fb70b5d7ceee19"><area href="/wiki/Progetto:Forme_di_vita/Come_leggere_il_tassobox" shape="rect" coords="0,0,50,50" alt="Progetto:Forme di vita/Come leggere il tassobox" title="Progetto:Forme di vita/Come leggere il tassobox" /></map><figcaption>Come leggere il tassobox</figcaption></figure></div>Saccharomyces carlsbergensis</th></tr><tr><td class="sinottico_testo_centrale" colspan="2"><span class="skin-invert itwiki-preserve-color" typeof="mw:File"><a href="/wiki/File:Fungi_missing.png" class="mw-file-description" title="Immagine di Saccharomyces carlsbergensis mancante"><img alt="Immagine di Saccharomyces carlsbergensis mancante" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Fungi_missing.png/220px-Fungi_missing.png" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Fungi_missing.png/330px-Fungi_missing.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Fungi_missing.png/440px-Fungi_missing.png 2x" data-file-width="1000" data-file-height="667" /></a></span></td></tr><tr class="sinottico_divisione"><th colspan="2" style="background-color: lightblue; color: black;"><a href="/wiki/Classificazione_scientifica" title="Classificazione scientifica">Classificazione scientifica</a></th></tr><tr><th style="font-weight:normal"><a href="/wiki/Dominio_(biologia)" title="Dominio (biologia)">Dominio</a></th><td><a href="/wiki/Eukaryota" title="Eukaryota">Eukaryota</a> </td></tr><tr><th style="font-weight:normal; width:50%"><a href="/wiki/Regno_(biologia)" title="Regno (biologia)">Regno</a></th><td><a href="/wiki/Fungi" title="Fungi">Fungi</a> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Phylum" title="Phylum">Phylum</a></th><td><a href="/wiki/Ascomycota" title="Ascomycota">Ascomycota</a> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Subphylum" title="Subphylum">Sottodivisione</a></th><td><a href="/wiki/Saccharomycotina" class="mw-redirect" title="Saccharomycotina">Saccharomycotina</a> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Classe_(tassonomia)" title="Classe (tassonomia)">Classe</a></th><td><a href="/wiki/Saccharomycetes" class="mw-redirect" title="Saccharomycetes">Saccharomycetes</a> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Ordine_(tassonomia)" title="Ordine (tassonomia)">Ordine</a></th><td><a href="/wiki/Saccharomycetales" title="Saccharomycetales">Saccharomycetales</a> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Famiglia_(tassonomia)" title="Famiglia (tassonomia)">Famiglia</a></th><td><a href="/w/index.php?title=Saccharomycetaceae&action=edit&redlink=1" class="new" title="Saccharomycetaceae (la pagina non esiste)">Saccharomycetaceae</a> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Genere_(tassonomia)" title="Genere (tassonomia)">Genere</a></th><td><i><a href="/wiki/Saccharomyces" title="Saccharomyces">Saccharomyces</a></i> </td></tr><tr><th style="font-weight:normal"><a href="/wiki/Specie" title="Specie">Specie</a></th><td><i><b>S. carlsbergensis</b></i> </td></tr><tr class="sinottico_divisione"><th colspan="2" style="background-color: lightblue; color: black;"><a href="/wiki/Nomenclatura_binomiale" title="Nomenclatura binomiale">Nomenclatura binomiale</a></th></tr><tr><td class="sinottico_testo_centrale" colspan="2"><i><b>Saccharomyces carlsbergensis</b></i><br /><small>Nguyen & Gaillardin ex. <a href="/w/index.php?title=Martinus_Willem_Beijerink&action=edit&redlink=1" class="new" title="Martinus Willem Beijerink (la pagina non esiste)">Beijerinck</a></small> </td></tr><tr class="sinottico_divisione"><th colspan="2" style="background-color: lightblue; color: black;"><a href="/wiki/Sinonimo_(tassonomia)" title="Sinonimo (tassonomia)">Sinonimi</a></th></tr><tr><td class="sinottico_testo_centrale" colspan="2">  <p><i>Saccharomyces pastorianus</i> </p> </td></tr></tbody></table> <p><i><b>Saccharomyces carlsbergensis</b></i>, attualmente denominato più correttamente <i><b>Saccharomyces pastorianus</b></i>, è un <a href="/wiki/Lievito" title="Lievito">lievito</a> utilizzato per la produzione della <a href="/wiki/Birra" title="Birra">birra</a> denominata <i><a href="/wiki/Lager_(birra)" title="Lager (birra)">lager</a></i>, a bassa <a href="/wiki/Fermentazione_alcolica" title="Fermentazione alcolica">fermentazione</a>. Questo tipo di lievito è stato analizzato originariamente da <a href="/wiki/Emil_Christian_Hansen" title="Emil Christian Hansen">Emil Christian Hansen</a>, che lavorava per il birrificio danese <a href="/wiki/Carlsberg" title="Carlsberg">Carlsberg</a>, che gli ha dato il nome. </p><p>Si tratta di un <a href="/w/index.php?title=Ibrido_(biologia)&action=edit&redlink=1" class="new" title="Ibrido (biologia) (la pagina non esiste)">ibrido</a> fra <i><a href="/wiki/Saccharomyces_bayanus" title="Saccharomyces bayanus">Saccharomyces bayanus</a></i> e <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i>: come tale, presenta una buona quantità di similitudini sia genotipiche che fenotipiche con le due specie genitrici<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>, anche se il suo <a href="/wiki/Genoma" title="Genoma">genoma</a> è lungo circa una volta e mezzo quello di queste ultime, poiché include quasi per intero i genomi delle due specie ibridate<sup id="cite_ref-Montrocher_2-0" class="reference"><a href="#cite_note-Montrocher-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>. </p><p>Da <i>S. bayanus</i>, la specie eredita gran parte dei geni, oltre che il <a href="/wiki/DNA_mitocondriale" title="DNA mitocondriale">DNA mitocondriale</a> e quello <a href="/w/index.php?title=DNA_ribosomiale&action=edit&redlink=1" class="new" title="DNA ribosomiale (la pagina non esiste)">ribosomiale</a><sup id="cite_ref-Montrocher_2-1" class="reference"><a href="#cite_note-Montrocher-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>: questo comporta numerosi adattamenti nella crescita che la specie condivide con quest'ultimo, infatti come <i>S. bayanus</i> non cresce a temperature superiori ai 34 °C, mentre <i>S. cerevisiae</i> cresce anche a 40 °C, ed allo stesso modo <i>S. carlsbergensis</i> cresce meglio di <i>S. cerevisiae</i> al di sotto dei 12 °C </p><p>I geni LEU2, MET2 ed ACB1 di questa specie hanno mostrato un alto grado di affinità con quelli della specie <i><a href="/w/index.php?title=Saccharomyces_monacensis&action=edit&redlink=1" class="new" title="Saccharomyces monacensis (la pagina non esiste)">Saccharomyces monacensis</a></i>, il che ha indotto alcuni studiosi ad ipotizzare che si tratti di un ibrido di questa specie con <i>S. cerevisiae</i> anziché con <i>S. bayanus</i><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><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>, tuttavia ulteriori analisi hanno chiarito che tale affinità è data da uno stretto legame di parentela, piuttosto che da una effettiva discendenza di una specie dall'altra. </p> <div class="mw-heading mw-heading2"><h2 id="Note">Note</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Saccharomyces_carlsbergensis&veaction=edit&section=1" title="Modifica la sezione Note" class="mw-editsection-visualeditor"><span>modifica</span></a><span class="mw-editsection-divider"> | </span><a href="/w/index.php?title=Saccharomyces_carlsbergensis&action=edit&section=1" title="Edit section's source code: Note"><span>modifica wikitesto</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-1"><a href="#cite_ref-1"><b>^</b></a> <span class="reference-text">Tamai Y, Momma T, Yoshimoto H, Kaneko Y., <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/entrez/query.fcgi?holding=npg&cmd=Retrieve&db=PubMed&list_uids=9717238&dopt=Abstract">Co-existence of two types of chromosome in the bottom fermenting yeast, Saccharomyces pastorianus.</a>, Yeast. 1998 Jul;14(10):923-33.</span> </li> <li id="cite_note-Montrocher-2"><span class="mw-cite-backlink"><b>^</b> <sup><i><a href="#cite_ref-Montrocher_2-0">a</a></i></sup> <sup><i><a href="#cite_ref-Montrocher_2-1">b</a></i></sup></span> <span class="reference-text">Montrocher R <i>et al</i> (1998), <a rel="nofollow" class="external text" href="http://ijs.sgmjournals.org/cgi/reprint/48/1/295.pdf">Phylogenetic Analysis of the Saccharomyces cerevisiae Group Based on Polymorphisms of the rDNA Spacer Sequences</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20081217025938/http://ijs.sgmjournals.org/cgi/reprint/48/1/295.pdf">Archiviato</a> il 17 dicembre 2008 in <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>., Int. J. Syst. Bacteriol., vol 48, pp 295–303</span> </li> <li id="cite_note-3"><a href="#cite_ref-3"><b>^</b></a> <span class="reference-text">Piskur, J., S. Smole, C. Groth, R.F. Petersen and M.B. Pedersen (1998), <a rel="nofollow" class="external text" href="http://ijs.sgmjournals.org/cgi/content/abstract/48/3/1015">Structure and genetic stability of mitochondrial genomes vary among yeasts of the genus Saccharomyces.</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20080423010351/http://ijs.sgmjournals.org/cgi/content/abstract/48/3/1015">Archiviato</a> il 23 aprile 2008 in <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>. Int. J. System. Bacteriol. 48: 1015-1024.</span> </li> <li id="cite_note-4"><a href="#cite_ref-4"><b>^</b></a> <span class="reference-text">Hansen J and Kielland-Brandt MC (1994), <a rel="nofollow" class="external text" href="https://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6T39-47PNX4N-B3&_coverDate=12%2F31%2F1994&_alid=512875340&_rdoc=1&_fmt=&_orig=search&_qd=1&_cdi=4941&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=ee8b387b7dd2fe32888a8cc8272d31d6">Saccharomyces carlsbergensis contains two functional MET2 alleles similar to homologues from S. cerevisiae and S. monacensis.</a> Gene 140(1):33-40</span> </li> <li id="cite_note-5"><a href="#cite_ref-5"><b>^</b></a> <span class="reference-text">Casey GP, Pedersen MB (1988), <a rel="nofollow" class="external text" href="http://db.yeastgenome.org/cgi-bin/reference/reference.pl?dbid=S000078250">DNA sequence polymorphisms in the genus Saccharomyces. V. Cloning and characterization of a LEU2 gene from S. carlsbergensis.</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20071023121139/http://db.yeastgenome.org/cgi-bin/reference/reference.pl?dbid=S000078250">Archiviato</a> il 23 ottobre 2007 in <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>. Carlsberg Res Commun 53(3):209-19</span> </li> <li id="cite_note-6"><a href="#cite_ref-6"><b>^</b></a> <span class="reference-text">Borsting C, et al. (1997),<a rel="nofollow" class="external text" href="http://db.yeastgenome.org/cgi-bin/reference/reference.pl?dbid=S000040754">Saccharomyces carlsbergensis contains two functional genes encoding the acyl-CoA binding protein, one similar to the ACB1 gene from S. cerevisiae and one identical to the ACB1 gene from S. monacensis.</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20071023120018/http://db.yeastgenome.org/cgi-bin/reference/reference.pl?dbid=S000040754">Archiviato</a> il 23 ottobre 2007 in <a href="/wiki/Internet_Archive" title="Internet Archive">Internet Archive</a>. Yeast 13(15):1409-21</span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Collegamenti_esterni">Collegamenti esterni</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Saccharomyces_carlsbergensis&veaction=edit&section=2" title="Modifica la sezione Collegamenti esterni" class="mw-editsection-visualeditor"><span>modifica</span></a><span class="mw-editsection-divider"> | </span><a href="/w/index.php?title=Saccharomyces_carlsbergensis&action=edit&section=2" title="Edit section's source code: Collegamenti esterni"><span>modifica wikitesto</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li class="mw-empty-elt"></li> <li><cite id="CITEREFBritannica.com" class="citation web" style="font-style:normal">(<span style="font-weight:bolder; font-size:80%"><abbr title="inglese">EN</abbr></span>) <a rel="nofollow" class="external text" href="https://www.britannica.com/science/Saccharomyces-carlsbergensis"><span style="font-style:italic;">Saccharomyces carlsbergensis</span></a>, su <span style="font-style:italic;"><a href="/wiki/Enciclopedia_Britannica" title="Enciclopedia Britannica">Enciclopedia Britannica</a></span>, Encyclopædia Britannica, Inc.</cite> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q20747163#P1417" title="Modifica su Wikidata"><img alt="Modifica su Wikidata" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/73/Blue_pencil.svg/10px-Blue_pencil.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/73/Blue_pencil.svg/15px-Blue_pencil.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/73/Blue_pencil.svg/20px-Blue_pencil.svg.png 2x" data-file-width="600" data-file-height="600" /></a></span></li></ul> <style data-mw-deduplicate="TemplateStyles:r141815314">.mw-parser-output .navbox{border:1px solid #aaa;clear:both;margin:auto;padding:2px;width:100%}.mw-parser-output .navbox th{padding-left:1em;padding-right:1em;text-align:center}.mw-parser-output .navbox>tbody>tr:first-child>th{background:#ccf;font-size:90%;width:100%;color:var(--color-base,black)}.mw-parser-output .navbox_navbar{float:left;margin:0;padding:0 10px 0 0;text-align:left;width:6em}.mw-parser-output .navbox_title{font-size:110%}.mw-parser-output .navbox_abovebelow{background:#ddf;font-size:90%;font-weight:normal}.mw-parser-output .navbox_group{background:#ddf;font-size:90%;padding:0 10px;white-space:nowrap}.mw-parser-output .navbox_list{font-size:90%;width:100%}.mw-parser-output .navbox_list a{white-space:nowrap}html:not(.vector-feature-night-mode-enabled) .mw-parser-output .navbox_odd{background:#fdfdfd;color:var(--color-base,black)}html:not(.vector-feature-night-mode-enabled) .mw-parser-output .navbox_even{background:#f7f7f7;color:var(--color-base,black)}.mw-parser-output .navbox a.mw-selflink{color:var(--color-base,black)}.mw-parser-output .navbox_center{text-align:center}.mw-parser-output .navbox .navbox_image{padding-left:7px;vertical-align:middle;width:0}.mw-parser-output .navbox+.navbox{margin-top:-1px}.mw-parser-output .navbox .mw-collapsible-toggle{font-weight:normal;text-align:right;width:7em}body.skin--responsive .mw-parser-output .navbox_image img{max-width:none!important}.mw-parser-output .subnavbox{margin:-3px;width:100%}.mw-parser-output .subnavbox_group{background:#e6e6ff;padding:0 10px}@media screen{html.skin-theme-clientpref-night .mw-parser-output .navbox>tbody>tr:first-child>th{background:var(--background-color-interactive)!important}html.skin-theme-clientpref-night .mw-parser-output .navbox th{color:var(--color-base)!important}html.skin-theme-clientpref-night .mw-parser-output .navbox_abovebelow,html.skin-theme-clientpref-night .mw-parser-output .navbox_group{background:var(--background-color-interactive-subtle)!important}html.skin-theme-clientpref-night .mw-parser-output .subnavbox_group{background:var(--background-color-neutral-subtle)!important}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbox>tbody>tr:first-child>th{background:var(--background-color-interactive)!important}html.skin-theme-clientpref-os .mw-parser-output .navbox th{color:var(--color-base)!important}html.skin-theme-clientpref-os .mw-parser-output .navbox_abovebelow,html.skin-theme-clientpref-os .mw-parser-output .navbox_group{background:var(--background-color-interactive-subtle)!important}html.skin-theme-clientpref-os .mw-parser-output .subnavbox_group{background:var(--background-color-neutral-subtle)!important}}</style><table class="navbox mw-collapsible mw-collapsed noprint metadata" id="navbox-Organismi_modello"><tbody><tr><th colspan="2"><div class="navbox_navbar"><div class="noprint plainlinks" style="background-color:transparent; 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