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<h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Chromatin</span></h1>
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<table class="infobox hintergrundfarbe-basis float-right" style="margin:0 0 1em 1.5em; border: 1px solid #CCD2D9; width: 200px; text-align: center; border-collapse: collapse; font-size: 90%;">

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<th style="background:#ABCDEF; color:#202122; border: 1px solid #ABCDEF;">Übergeordnet
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<td style="padding: 0 .5em;"><a href="Chromosom" title="Chromosom">Chromosom</a>
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<th style="background:#ABCDEF; color:#202122; border: 1px solid #ABCDEF;">Untergeordnet
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<td style="padding: 0 .5em;"><a href="Euchromatin" title="Euchromatin">Euchromatin</a><br><a href="Heterochromatin" title="Heterochromatin">Heterochromatin</a><br>zytoplasm./nukl. Chromatin<br>aktives/ruhendes Chromatin
</td></tr>
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<th style="background:#ABCDEF; color:ä202122; border: 1px solid #ABCDEF;"><a href="Gene_Ontology" title="Gene Ontology">Gene Ontology</a>
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<td style="text-align:left; padding: 0 .5em;"><a rel="nofollow" class="external text" href="http://www.ebi.ac.uk/QuickGO/GTerm?id=GO:0000785">QuickGO</a>
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<p><b>Chromatin</b> ist das Material, aus dem die <a href="Chromosom" title="Chromosom">Chromosomen</a> bestehen. Es handelt sich um einen Komplex aus <a href="Desoxyribonukleins%C3%A4ure" title="Desoxyribonukleinsäure">DNA</a> und speziellen <a href="Protein" title="Protein">Proteinen</a>, von denen wiederum etwa die Hälfte <a href="Histon" title="Histon">Histone</a> sind. Der Name kommt von griech. <i>chroma</i> (Farbe), weil sich Chromatin mit basischen Kernfarbstoffen anfärben lässt. Im Lichtmikroskop erscheint es als sichtbares Fadengerüst im Zellkern einer <a href="Eukaryoten" title="Eukaryoten">eukaryotischen</a> <a href="Zelle_(Biologie)" title="Zelle (Biologie)">Zelle</a>. Im funktionalen Sinn gilt alles, was sich während der Teilung des <a href="Zellkern" title="Zellkern">Zellkerns</a> (<i><a href="Mitose" title="Mitose">Mitose</a></i> oder <i><a href="Meiose" title="Meiose">Meiose</a></i>) in den Chromosomen wiederfindet, als Chromatin – ausgenommen einige <a href="Strukturprotein" title="Strukturprotein">Strukturproteine</a>. Chromatin ist neben den <a href="Nucleolus" title="Nucleolus">Nucleoli</a>, der <a href="Karyoplasma" title="Karyoplasma">Kern-Grundsubstanz</a> und der <a href="Kernh%C3%BClle" title="Kernhülle">Kernhülle</a> eine wichtige Strukturkomponente des Zellkerns (Nucleus).<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>
</p><p>Chromatin besteht aus der DNA, die um die Histone gewickelt ist, sowie aus weiteren Proteinen, die sich an die DNA anlagern. DNA und Histone bilden die <a href="Nucleosom" class="mw-redirect" title="Nucleosom">Nucleosomen</a>, die kettenförmig aneinandergereiht sind. Die Nucleosomen werden mit Hilfe der Nichthiston-Proteine dichter gepackt. Chromatin ist somit das Produkt von Interaktionen der eukaryotischen DNA mit unterschiedlichen DNA-Bindeproteinen, die einen kompakten filamentösen Komplex bilden, den sogenannten Desoxyribonucleoprotein-Komplex, man spricht auch von Chromatinfasern oder Chromatinfäden (englisch: <i>chromatin fibers</i>). Durch die Komplexbildung werden die langen chromosomalen DNA-Stränge in ihrer Länge um das rund 10.000- bis 50.000-fache verkürzt (kondensiert), sodass sie in den Zellkern passen. Trotz der dichten Packung der DNA liegen die Chromosomen weiterhin in einer Form vor, die regulatorischen Proteinen Zugang zur DNA erlaubt, so dass die Biosynthese von RNA und Proteinen aus den genetischen Informationen (<a href="Genexpression" title="Genexpression">Genexpression</a>) bzw. die Duplikation der chromosomalen DNA (<a href="Replikation" title="Replikation">Replikation</a>) möglich ist.<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>
</p><p>Während der Mitose und Meiose <a href="Kondensation_(Genetik)" class="mw-redirect" title="Kondensation (Genetik)">kondensieren</a> die Chromosomen, so dass sie im Lichtmikroskop erkennbar werden. Die kleinsten lichtmikroskopisch sichtbaren Chromatinstrukturen nennt man <a href="Chromonema" title="Chromonema">Chromonema</a>.
</p><p>Das Verständnis der Chromatinstruktur und ihres Beitrags zu Regulation der <a href="Gen" title="Gen">Gene</a> ist Gegenstand der <a href="Epigenetik" title="Epigenetik">Epigenetik</a>.
</p><p>Chromatinstrukturen machen <a href="St%C3%A4bchen_(Auge)" title="Stäbchen (Auge)">Stäbchen</a> bei nachtaktiven Säugetieren empfindlicher, da sie die Lichtausbreitung beeinflussen. Bei Nicht-Säugern ist das Phänomen noch nicht untersucht worden (Stand 2010).<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>
</p>

<div class="mw-heading mw-heading2"><h2 id="Chromatin-Typen">Chromatin-Typen</h2></div>
<p>Es werden zwei Typen von Chromatin unterschieden:
</p>
<ul><li><a href="Euchromatin" title="Euchromatin">Euchromatin</a>, dessen DNA aktiv ist, d.&nbsp;h., zu Proteinen <a href="Genexpression" title="Genexpression">exprimiert</a> werden kann. Die euchromatischen Abschnitte des Chromosoms weisen keine Unterschiede in ihrer Struktur auf, gleichgültig, in welchem Kondensationsgrad sich ein Chromosom befindet.</li>
<li><a href="Heterochromatin" title="Heterochromatin">Heterochromatin</a>, das hauptsächlich aus inaktiver DNA besteht. Es scheint strukturelle Funktionen in den verschiedenen Kondensationsstufen auszuüben. Die heterochromatischen Abschnitte des Chromosoms weisen in der <a href="Interphase" title="Interphase">Interphase</a> den gleichen Kondensationsgrad auf wie in der <a href="Metaphase" class="mw-redirect" title="Metaphase">Metaphase</a>, d.&nbsp;h., es bleibt auch im Interphasekern kondensiert und tritt in Form dichter <a href="Chromozentren" class="mw-redirect" title="Chromozentren">Chromozentren</a> in Erscheinung. Heterochromatin kann in zwei Untertypen unterteilt werden:
<ul><li>Konstitutives Heterochromatin, das nie exprimiert wird. Es findet sich im Bereich des Centromers und besteht gewöhnlich aus repetitiven (sich wiederholenden) DNA-Sequenzen.</li>
<li>Fakultatives Heterochromatin, das manchmal exprimiert wird.</li></ul></li></ul>
<p>Eine weitere begriffliche Abgrenzung kann somit auch nach den Kernteilungsphasen getroffen werden: Hierbei ist das <i>Interphasechromatin</i> gegenüber dem <i>Metaphasechromatin</i> mit seinen sehr kompakten Chromosomen stark aufgelockert.
</p><p>Prokaryonten haben im Gegensatz zu Eukaryonten eine ringförmige DNA-Struktur. Die Eukaryonten haben Chromosomen, die die Struktur der DNA bilden.
</p>
<div class="mw-heading mw-heading2"><h2 id="Zeittafel_wichtiger_Entdeckungen">Zeittafel wichtiger Entdeckungen</h2></div>

<ul><li>1842: <a href="Chromosom" title="Chromosom">Chromosom</a> (Struktur) (<a href="Carl_Wilhelm_von_N%C3%A4geli" title="Carl Wilhelm von Nägeli">Carl Wilhelm von Nägeli</a>)</li>
<li>1874: <a href="Nukleins%C3%A4ure" class="mw-redirect" title="Nukleinsäure">Nukleinsäure</a> (<a href="Friedrich_Miescher" title="Friedrich Miescher">Friedrich Miescher</a>)</li>
<li>1879: prägt <a href="Walther_Flemming" title="Walther Flemming">Walther Flemming</a> den Begriff Chromatin.</li>
<li>1883: <a href="August_Weismann" title="August Weismann">August Weismann</a> verbindet Chromatin mit der Vererbung.</li>
<li>1884: <a href="Albrecht_Kossel" title="Albrecht Kossel">Albrecht Kossel</a> entdeckt <a href="Histon" title="Histon">Histone</a>.</li>
<li>1888: <a href="Walter_Sutton" title="Walter Sutton">Sutton</a> und <a href="Theodor_Boveri_(Biologe)" class="mw-redirect" title="Theodor Boveri (Biologe)">Boveri</a> schlagen die Theorie der Kontinuität von Chromatin während des Zellzyklus vor.<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></li>
<li>1888: <a href="Heinrich_Wilhelm_Waldeyer" class="mw-redirect" title="Heinrich Wilhelm Waldeyer">Wilhelm Waldeyer</a> prägt den Begriff <a href="Chromosom" title="Chromosom">Chromosom</a></li>
<li>1910: Chromosomen sind die Träger der Gene (<a href="Thomas_Hunt_Morgan" title="Thomas Hunt Morgan">Thomas Hunt Morgan</a>)</li>
<li>1928: <a href="Emil_Heitz_(Botaniker)" title="Emil Heitz (Botaniker)">Emil Heitz</a> prägt den Begriff <a href="Heterochromatin" title="Heterochromatin">Heterochromatin</a> und <a href="Euchromatin" title="Euchromatin">Euchromatin</a>.</li>
<li>1942: <a href="Conrad_Hal_Waddington" title="Conrad Hal Waddington">Conrad Waddington</a> postuliert die <a href="Epigenetik" title="Epigenetik">epigenetischen Landschaften</a>.</li>
<li>ca. 1945 Basenpaarung von <a href="Adenin" title="Adenin">Adenin</a> und <a href="Thymin" title="Thymin">Thymin</a> sowie <a href="Cytosin" title="Cytosin">Cytosin</a> und <a href="Guanin" title="Guanin">Guanin</a> postuliert (<a href="Erwin_Chargaff" title="Erwin Chargaff">Erwin Chargaff</a>, <a href="Chargaff-Regeln" title="Chargaff-Regeln">Chargaff-Regeln</a>)</li>
<li>1948: Rollin Hotchkiss entdeckt <a href="DNA-Methylierung" title="DNA-Methylierung">DNA-Methylierung</a><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></li>
<li>1953: Die DNA-Struktur wird aufgeklärt von <a href="James_Watson" title="James Watson">James Watson</a>, <a href="Francis_Crick" title="Francis Crick">Francis Crick</a>, <a href="Maurice_Wilkins" title="Maurice Wilkins">Maurice Wilkins</a>, <a href="Rosalind_Franklin" title="Rosalind Franklin">Rosalind Franklin</a><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></li>
<li>1961: Aufklärung der genetischen Struktur (<a href="Marshall_Warren_Nirenberg" title="Marshall Warren Nirenberg">Marshall Warren Nirenberg</a>, <a href="Heinrich_Matthaei" title="Heinrich Matthaei">Heinrich Matthaei</a>)</li>
<li>1961: <a href="Mary_Frances_Lyon" title="Mary Frances Lyon">Mary Lyon</a> postuliert das Prinzip der <a href="X-Inaktivierung" title="X-Inaktivierung">X-Inaktivierung</a>.</li>
<li>1966: <a href="Epigenetik" title="Epigenetik">Histon-Modifikationen</a>/-Acetylierung (Vincent Allfrey)</li>
<li>1973:/1974 Chromatinfasern werden entdeckt</li>
<li>1973:–75 Vom nu-Body zum <a href="Nukleosom" title="Nukleosom">Nukleosom</a> (Ada Olins, Donald Olins, <a href="Roger_Kornberg" class="mw-redirect" title="Roger Kornberg">Roger Kornberg</a>)</li>
<li>1975: Nukleosomen-Überstruktur/<a href="Solenoidstruktur" title="Solenoidstruktur">Solenoid</a> (John T. Finch und <a href="Aaron_Klug" title="Aaron Klug">Aaron Klug</a>)</li>
<li>1975: <a href="Pierre_Chambon" title="Pierre Chambon">Pierre Chambon</a> prägt den Begriff der <a href="Nukleosom" title="Nukleosom">Nukleosomen</a>.</li>
<li>1976: Chromatinfäden werden entdeckt<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></li>
<li>1982: <a href="Chromosomenterritorium" title="Chromosomenterritorium">Chromosomenterritorien</a> werden entdeckt<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></li>
<li>1984: John T. Lis entwickelt die <a href="Chromatin-Immunpr%C3%A4zipitation" title="Chromatin-Immunpräzipitation">Chromatin-Immunpräzipitationstechnik</a>.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup></li>
<li>1996 wurde mit der <i><a href="Backhefe" title="Backhefe">Backhefe</a></i> das erste Genom eines Eukaryonten <a href="Liste_von_sequenzierten_Genomen" title="Liste von sequenzierten Genomen">veröffentlicht</a>.<sup id="cite_ref-Goffeau_10-0" class="reference"><a href="#cite_note-Goffeau-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup></li>
<li>2002: Job Dekker entwickelt die <a href="Chromosome_conformation_capture" title="Chromosome conformation capture">Chromosome Conformation Capture</a> (3C) Technik.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup></li>
<li>2006: Marieke Simons entwickelt die 4C-<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup>, Dostie die 5C-Methode<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup></li>
<li>2007: B. Franklin Pugh entwickelt die <a href="ChIP-Seq" title="ChIP-Seq">ChIP-Seq</a>-Technik.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li>
<li>2009: Lieberman-Aiden und Job Dekker erfindet die <a href="Chromosome_conformation_capture" title="Chromosome conformation capture">Hi-C-Technik</a><sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>, Melissa J. Fullwood erfindet die ChIA-Pet-Technik.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></li>
<li>2012: Eine Gruppe der Ren-Labs und die von Edith Heard und Job Dekker geleiteten Gruppen entdecken <a href="Topologically_associating_domain" title="Topologically associating domain">Topologically Associated Domains</a> (TADs) bei Säugetieren.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Literatur">Literatur</h2></div>
<p>Evolution:
</p>
<ul><li>R. Ammar, D. Torti u.&nbsp;a.: <i>Chromatin is an ancient innovation conserved between Archaea and Eukarya.</i> In: <i>eLife.</i> 1, 2012, S.&nbsp;e00078–e00078, <a href="https://doi.org/10.7554/eLife.00078" class="extiw external" title="doi:10.7554/eLife.00078">doi:10.7554/eLife.00078</a>.</li>
<li>Xavier Grau-Bové, Cristina Navarrete, Cristina Chiva, Thomas Pribasnig, Meritxell Antó, Guifré Torruella, Luis Javier Galindo, Bernd Franz Lang, David Moreira, Purificación López-Garcia, Iñaki Ruiz-Trillo, Christa Schleper, Eduard Sabidó, Arnau Sebé-Pedrós: <a rel="nofollow" class="external text" href="https://www.nature.com/articles/s41559-022-01771-6">A phylogenetic and proteomic reconstruction of eukaryotic chromatin evolution</a>. In: <i>Nature Ecology &amp; Evolution</i>, 9. Juni 2022, Band 6, S.&nbsp;1007–1023; <a href="https://doi.org/10.1038/s41559-022-01771-6" class="extiw external" title="doi:10.1038/s41559-022-01771-6">doi:10.1038/s41559-022-01771-6</a>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/35680998?dopt=Abstract">PMID 35680998</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7613034/">PMC&nbsp;7613034</a> (freier Volltext). Dazu:
<ul><li><a rel="nofollow" class="external text" href="https://media.springernature.com/full/springer-static/image/art%3A10.1038%2Fs41559-022-01771-6/MediaObjects/41559_2022_1771_Fig1_HTML.png">Fig.&nbsp;1: Diversity of post-translational modifications in eukaryotic canonical and variant histones</a>.</li>
<li><a rel="nofollow" class="external text" href="https://media.springernature.com/full/springer-static/image/art%3A10.1038%2Fs41559-022-01771-6/MediaObjects/41559_2022_1771_Fig3_HTML.png">Fig.&nbsp;3: Taxonomic distribution of chromatin-associated gene classes</a>.</li>
<li><a rel="nofollow" class="external text" href="https://media.springernature.com/full/springer-static/image/art%3A10.1038%2Fs41559-022-01771-6/MediaObjects/41559_2022_1771_Fig6_HTML.png">Fig.&nbsp;6: Chromatin evolution and eukaryogenesis</a>.</li>
<li>Xavier Grau-Bové: <a rel="nofollow" class="external text" href="https://github.com/sebepedroslab/chromatin-evolution-analysis">Repository: Chromatin evolution.</a> <a href="GitHub" title="GitHub">GitHub</a>.</li>
<li><a rel="nofollow" class="external text" href="https://www.sci.news/biology/eukaryotic-chromatin-evolution-10894.html">Chromatin First Evolved in Ancient Microbes 1–2 Billion Years Ago, New Research Suggests.</a> sci.news, 13. Juni 2022.</li>
<li><a rel="nofollow" class="external text" href="https://scitechdaily.com/shrouded-in-mystery-scientists-finally-discover-the-origin-of-chromatin/"><i>Shrouded in Mystery: Scientists Finally Discover the Origin of Chromatin</i>.</a> SciTechDaily, 24. August 2022.</li>
<li><a rel="nofollow" class="external text" href="https://www.sciencedaily.com/releases/2022/06/220609131933.htm"><i>Chromatin originated in ancient microbes one to two billion years ago</i>.</a> <i>Genomic and proteomic analysis reveals that the regulatory role of chromatin is a eukaryotic innovation</i>. ScienceDaily, 9. Juni 2022. Quelle: Center for Genomic Regulation (CRG).</li>
<li><a rel="nofollow" class="external text" href="https://www.crg.eu/en/news/chromatin-originated-ancient-microbes-one-two-billion-years-ago">Chromatin originated in ancient microbes one to two billion years ago.</a> Center for Genomic Regulation (CRG), 9. Juni 2022.</li></ul></li></ul>
<p>Histon-Modifikationen:
</p>
<ul><li>V. G. Allfrey: <cite style="font-style:italic">Structural modifications of histones and their possible role in the regulation of ribonucleic acid synthesis</cite>. In: <cite style="font-style:italic">Proceedings. Canadian Cancer Conference</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>6</span>, 1966, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>313–335</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/5934780?dopt=Abstract">PMID 5934780</a>.<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Structural+modifications+of+histones+and+their+possible+role+in+the+regulation+of+ribonucleic+acid+synthesis&amp;rft.au=V.+G.+Allfrey&amp;rft.btitle=Proceedings.+Canadian+Cancer+Conference&amp;rft.date=1966&amp;rft.genre=book&amp;rft.pages=313-335&amp;rft.pmid=5934780&amp;rft.volume=6" style="display:none">&nbsp;</span></li>
<li>B. G. Pogo, A. O. Pogo, V. G. Allfrey, A. E. Mirsky: <cite style="font-style:italic">Changing patterns of histone acetylation and RNA synthesis in regeneration of the liver.</cite> In: <cite style="font-style:italic"><a href="Proceedings_of_the_National_Academy_of_Sciences_of_the_United_States_of_America" title="Proceedings of the National Academy of Sciences of the United States of America">Proceedings of the National Academy of Sciences of the United States of America</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>59</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>4</span>, 1968, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>1337–1344</span>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC224872/">PMC&nbsp;224872</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Changing+patterns+of+histone+acetylation+and+RNA+synthesis+in+regeneration+of+the+liver.&amp;rft.au=B.+G.+Pogo%2C+A.+O.+Pogo%2C+V.+G.+Allfrey%2C+...&amp;rft.date=1968&amp;rft.genre=journal&amp;rft.issue=4&amp;rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&amp;rft.pages=1337-1344&amp;rft.pmc=224872&amp;rft.volume=59" style="display:none">&nbsp;</span></li></ul>
<p>Nukleosomen:
</p>
<ul><li>A. L. Olins, D. E. Olins: <cite style="font-style:italic">Spheroid chromatin units (v bodies)</cite>. In: <cite style="font-style:italic"><a href="Science" title="Science">Science</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>183</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>4122</span>, 1974, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>330–332</span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/4128918?dopt=Abstract">PMID 4128918</a>.<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Spheroid+chromatin+units+%28v+bodies%29&amp;rft.au=A.+L.+Olins%2C+D.+E.+Olins&amp;rft.date=1974&amp;rft.genre=journal&amp;rft.issue=4122&amp;rft.jtitle=Science&amp;rft.pages=330-332&amp;rft.pmid=4128918&amp;rft.volume=183" style="display:none">&nbsp;</span></li></ul>
<p>Solenoid-Modell:
</p>
<ul><li>J. T. Finch, A. Klug: <cite style="font-style:italic">Solenoidal model for superstructure in chromatin.</cite> In: <cite style="font-style:italic"><a href="Proceedings_of_the_National_Academy_of_Sciences_of_the_United_States_of_America" title="Proceedings of the National Academy of Sciences of the United States of America">Proceedings of the National Academy of Sciences of the United States of America</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>73</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>6</span>, 1976, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>1897–1901</span>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC430414/">PMC&nbsp;430414</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Solenoidal+model+for+superstructure+in+chromatin.&amp;rft.au=J.+T.+Finch%2C+A.+Klug&amp;rft.date=1976&amp;rft.genre=journal&amp;rft.issue=6&amp;rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&amp;rft.pages=1897-1901&amp;rft.pmc=430414&amp;rft.volume=73" style="display:none">&nbsp;</span></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Weblinks">Weblinks</h2></div>
<div class="sisterproject" style="margin:0.1em 0 0 0;"><div class="noresize noviewer" style="display:inline-block; line-height:10px; min-width:1.6em; text-align:center;" aria-hidden="true" role="presentation"><span class="mw-default-size" typeof="mw:File"><span title="Commons"></span></span></div><b><span class=""><a class="external text" href="https://commons.wikimedia.org/wiki/Category:Chromatin?uselang=de"><span lang="en">Commons</span>: Chromatin</a></span></b>&nbsp;– Sammlung von Bildern, Videos und Audiodateien</div>
<div class="sisterproject" style="margin:0.1em 0 0 0;"><span class="noviewer" style="display:inline-block; line-height:10px; min-width:1.6em; text-align:center;" aria-hidden="true" role="presentation"><span class="mw-default-size" typeof="mw:File"><span title="Wiktionary"></span></span></span><b><a href="https://de.wiktionary.org/wiki/Chromatin" class="extiw external" title="wikt:Chromatin">Wiktionary: Chromatin</a></b>&nbsp;– Bedeutungserklärungen, Wortherkunft, Synonyme, Übersetzungen</div>
<ul><li><a rel="nofollow" class="external text" href="http://www.chromdb.org/">ChromDB – The Chromatin Database</a></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Einzelnachweise">Einzelnachweise</h2></div>
<ol class="references">
<li id="cite_note-1"><span class="mw-cite-backlink"><a href="#cite_ref-1">↑</a></span> <span class="reference-text">Hans Kleinig, Peter Sitte: <i>Zellbiologie. Ein Lehrbuch.</i> 3. Auflage. Gustav Fischer Verlag, 1992, S. 176.</span>
</li>
<li id="cite_note-2"><span class="mw-cite-backlink"><a href="#cite_ref-2">↑</a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r261891140">
/* start https://de.wikipedia.org/ */


.mw-parser-output .webarchiv-memento a{color:inherit}


/* end https://de.wikipedia.org/ */
</style><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090722035437/http://www.chromdb.org/beginners.html"><i>Chromatin and chromosome structure</i>.</a> (<span class="webarchiv-memento"><a href="Webarchivierung#Begrifflichkeiten" title="Webarchivierung">Memento</a></span> vom 22. Juli 2009 im <i><a href="Internet_Archive" title="Internet Archive">Internet Archive</a></i>) The Chromatin Database; abgerufen am 12. Juni 2009.</span>
</li>
<li id="cite_note-3"><span class="mw-cite-backlink"><a href="#cite_ref-3">↑</a></span> <span class="reference-text"><a rel="nofollow" class="external text" href="http://www.schattenblick.de/infopool/natur/biologie/nbfor597.html">Nachtsehen – Wenn jedes Lichtquant zählt.</a> schattenblick.de, 17. April 2009.</span>
</li>
<li id="cite_note-4"><span class="mw-cite-backlink"><a href="#cite_ref-4">↑</a></span> <span class="reference-text">L.A.-C.P. Martins: <cite style="font-style:italic">Did Sutton and Boveri propose the so-called Sutton-Boveri chromosome hypothesis?</cite> In: <cite style="font-style:italic">Genetics and Molecular Biology</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>22</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>2</span>, Juni 1999, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a>&nbsp;<span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%221415-4757%22&amp;key=cql">1415-4757</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>261–272</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/10.1590/S1415-47571999000200022">10.1590/S1415-47571999000200022</a></span> (<a rel="nofollow" class="external text" href="http://www.scielo.br/scielo.php?script=sci_arttext&amp;pid=S1415-47571999000200022&amp;lng=en&amp;tlng=en">scielo.br</a> [abgerufen am 22.&nbsp;Juli 2019]).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Did+Sutton+and+Boveri+propose+the+so-called+Sutton-Boveri+chromosome+hypothesis%3F&amp;rft.au=L.A.-C.P.+Martins&amp;rft.date=1999-06&amp;rft.doi=10.1590%2FS1415-47571999000200022&amp;rft.genre=journal&amp;rft.issn=1415-4757&amp;rft.issue=2&amp;rft.jtitle=Genetics+and+Molecular+Biology&amp;rft.pages=261-272&amp;rft.volume=22" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-5"><span class="mw-cite-backlink"><a href="#cite_ref-5">↑</a></span> <span class="reference-text">Haoyang Lu, Xinzhou Liu, Yulin Deng, Hong Qing: <cite style="font-style:italic">DNA methylation, a hand behind neurodegenerative diseases</cite>. In: <cite style="font-style:italic">Frontiers in Aging Neuroscience</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>5</span>, 2013, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a>&nbsp;<span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%221663-4365%22&amp;key=cql">1663-4365</a></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/10.3389/fnagi.2013.00085">10.3389/fnagi.2013.00085</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24367332?dopt=Abstract">PMID 24367332</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3851782/">PMC&nbsp;3851782</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=DNA+methylation%2C+a+hand+behind+neurodegenerative+diseases&amp;rft.au=Haoyang+Lu%2C+Xinzhou+Liu%2C+Yulin+Deng%2C+...&amp;rft.date=2013&amp;rft.doi=10.3389%2Ffnagi.2013.00085&amp;rft.genre=journal&amp;rft.issn=1663-4365&amp;rft.jtitle=Frontiers+in+Aging+Neuroscience&amp;rft.pmc=3851782&amp;rft.pmid=24367332&amp;rft.volume=5" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-6"><span class="mw-cite-backlink"><a href="#cite_ref-6">↑</a></span> <span class="reference-text"><span class="cite"><a rel="nofollow" class="external text" href="https://profiles.nlm.nih.gov/SC/Views/Exhibit/narrative/doublehelix.html"><i>The Francis Crick Papers: The Discovery of the Double Helix, 1951–1953.</i></a> In: <i>profiles.nlm.nih.gov.</i><span class="Abrufdatum"> Abgerufen am 22.&nbsp;Juli 2019</span>.</span><span style="display: none;" class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Adc&amp;rfr_id=info%3Asid%2Fde.wikipedia.org%3AChromatin&amp;rft.title=The+Francis+Crick+Papers%3A+The+Discovery+of+the+Double+Helix%2C+1951%E2%80%931953&amp;rft.description=The+Francis+Crick+Papers%3A+The+Discovery+of+the+Double+Helix%2C+1951%E2%80%931953&amp;rft.identifier=https%3A%2F%2Fprofiles.nlm.nih.gov%2FSC%2FViews%2FExhibit%2Fnarrative%2Fdoublehelix.html">&nbsp;</span></span>
</li>
<li id="cite_note-7"><span class="mw-cite-backlink"><a href="#cite_ref-7">↑</a></span> <span class="reference-text">Ute Deichmann: <cite style="font-style:italic">Epigenetics: The origins and evolution of a fashionable topic</cite>. In: <cite style="font-style:italic">Developmental Biology</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>416</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>1</span>, August 2016, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>249–254</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/j.ydbio.2016.06.005">10.1016/j.ydbio.2016.06.005</a></span> (<a rel="nofollow" class="external text" href="https://linkinghub.elsevier.com/retrieve/pii/S0012160616302974">elsevier.com</a> [abgerufen am 22.&nbsp;Juli 2019]).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Epigenetics%3A+The+origins+and+evolution+of+a+fashionable+topic&amp;rft.au=Ute+Deichmann&amp;rft.date=2016-08&amp;rft.doi=10.1016%2Fj.ydbio.2016.06.005&amp;rft.genre=journal&amp;rft.issue=1&amp;rft.jtitle=Developmental+Biology&amp;rft.pages=249-254&amp;rft.volume=416" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-8"><span class="mw-cite-backlink"><a href="#cite_ref-8">↑</a></span> <span class="reference-text">T. Cremer, M. Cremer: <cite style="font-style:italic">Chromosome Territories</cite>. In: <cite style="font-style:italic">Cold Spring Harbor Perspectives in Biology</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>2</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>3</span>, 1.&nbsp;März 2010, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a>&nbsp;<span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%221943-0264%22&amp;key=cql">1943-0264</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>a003889–a003889</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/10.1101/cshperspect.a003889">10.1101/cshperspect.a003889</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/20300217?dopt=Abstract">PMID 20300217</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2829961/">PMC&nbsp;2829961</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Chromosome+Territories&amp;rft.au=T.+Cremer%2C+M.+Cremer&amp;rft.date=2010-03-01&amp;rft.doi=10.1101%2Fcshperspect.a003889&amp;rft.genre=journal&amp;rft.issn=1943-0264&amp;rft.issue=3&amp;rft.jtitle=Cold+Spring+Harbor+Perspectives+in+Biology&amp;rft.pages=a003889-a003889&amp;rft.pmc=2829961&amp;rft.pmid=20300217&amp;rft.volume=2" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-9"><span class="mw-cite-backlink"><a href="#cite_ref-9">↑</a></span> <span class="reference-text">D. S. Gilmour, J. T. Lis: <cite style="font-style:italic">Detecting protein-DNA interactions in vivo: distribution of RNA polymerase on specific bacterial genes.</cite> In: <cite style="font-style:italic">Proceedings of the National Academy of Sciences</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>81</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>14</span>, 1.&nbsp;Juli 1984, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a>&nbsp;<span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%220027-8424%22&amp;key=cql">0027-8424</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>4275–4279</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.81.14.4275">10.1073/pnas.81.14.4275</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/6379641?dopt=Abstract">PMID 6379641</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC345570/">PMC&nbsp;345570</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Detecting+protein-DNA+interactions+in+vivo%3A+distribution+of+RNA+polymerase+on+specific+bacterial+genes.&amp;rft.au=D.+S.+Gilmour%2C+J.+T.+Lis&amp;rft.date=1984-07-01&amp;rft.doi=10.1073%2Fpnas.81.14.4275&amp;rft.genre=journal&amp;rft.issn=0027-8424&amp;rft.issue=14&amp;rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences&amp;rft.pages=4275-4279&amp;rft.pmc=345570&amp;rft.pmid=6379641&amp;rft.volume=81" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-Goffeau-10"><span class="mw-cite-backlink"><a href="#cite_ref-Goffeau_10-0">↑</a></span> <span class="reference-text">A Goffeau, BG Barrell, H Bussey, R. W. Davis, B. Dujon, H. Feldmann, F. Galibert, J. D. Hoheisel, C. Jacq, M. Johnston, E. J. Louis, H. W. Mewes, Y. Murakami, P. Philippsen, H. Tettelin, S. G. Oliver: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Life with 6000 genes</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic"><a href="Science" title="Science">Science</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>274</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>5287</span>, Oktober 1996, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>546,&nbsp;563–567</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/10.1126/science.274.5287.546">10.1126/science.274.5287.546</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/8849441?dopt=Abstract">PMID 8849441</a>, <a href="Bibcode" title="Bibcode">bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1996Sci...274..546G">1996Sci...274..546G</a> (englisch).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Life+with+6000+genes&amp;rft.au=A+Goffeau%2C+BG+Barrell%2C+H+Bussey%2C+...&amp;rft.date=1996-10&amp;rft.doi=10.1126%2Fscience.274.5287.546&amp;rft.genre=journal&amp;rft.issue=5287&amp;rft.jtitle=Science&amp;rft.pages=546%2C+563-567&amp;rft.pmid=8849441&amp;rft.volume=274" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-11"><span class="mw-cite-backlink"><a href="#cite_ref-11">↑</a></span> <span class="reference-text">J. Dekker: <cite style="font-style:italic">Capturing Chromosome Conformation</cite>. In: <cite style="font-style:italic"><a href="Science" title="Science">Science</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>295</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>5558</span>, 15.&nbsp;Februar 2002, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>1306–1311</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/10.1126/science.1067799">10.1126/science.1067799</a></span>.<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Capturing+Chromosome+Conformation&amp;rft.au=J.+Dekker&amp;rft.date=2002-02-15&amp;rft.doi=10.1126%2Fscience.1067799&amp;rft.genre=journal&amp;rft.issue=5558&amp;rft.jtitle=Science&amp;rft.pages=1306-1311&amp;rft.volume=295" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-12"><span class="mw-cite-backlink"><a href="#cite_ref-12">↑</a></span> <span class="reference-text">Marieke Simonis, Petra Klous, Erik Splinter, Yuri Moshkin, Rob Willemsen: <cite style="font-style:italic">Nuclear organization of active and inactive chromatin domains uncovered by chromosome conformation capture–on-chip (4C)</cite>. In: <cite style="font-style:italic">Nature Genetics</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>38</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>11</span>, November 2006, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a>&nbsp;<span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%221061-4036%22&amp;key=cql">1061-4036</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>1348–1354</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/10.1038/ng1896">10.1038/ng1896</a></span>.<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Nuclear+organization+of+active+and+inactive+chromatin+domains+uncovered+by+chromosome+conformation+capture-on-chip+%284C%29&amp;rft.au=Marieke+Simonis%2C+Petra+Klous%2C+Erik+Splinter%2C+...&amp;rft.date=2006-11&amp;rft.doi=10.1038%2Fng1896&amp;rft.genre=journal&amp;rft.issn=1061-4036&amp;rft.issue=11&amp;rft.jtitle=Nature+Genetics&amp;rft.pages=1348-1354&amp;rft.volume=38" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-13"><span class="mw-cite-backlink"><a href="#cite_ref-13">↑</a></span> <span class="reference-text">J. Dostie, T. A. Richmond, R. A. Arnaout, R. R. Selzer, W. L. Lee: <cite style="font-style:italic">Chromosome Conformation Capture Carbon Copy (5C): A massively parallel solution for mapping interactions between genomic elements</cite>. In: <cite style="font-style:italic">Genome Research</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>16</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>10</span>, 1.&nbsp;Oktober 2006, <a href="Internationale_Standardnummer_f%C3%BCr_fortlaufende_Sammelwerke" title="Internationale Standardnummer für fortlaufende Sammelwerke">ISSN</a>&nbsp;<span style="white-space:nowrap"><a rel="nofollow" class="external text" href="https://zdb-katalog.de/list.xhtml?t=iss%3D%221088-9051%22&amp;key=cql">1088-9051</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>1299–1309</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/10.1101/gr.5571506">10.1101/gr.5571506</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/16954542?dopt=Abstract">PMID 16954542</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1581439/">PMC&nbsp;1581439</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Chromosome+Conformation+Capture+Carbon+Copy+%285C%29%3A+A+massively+parallel+solution+for+mapping+interactions+between+genomic+elements&amp;rft.au=J.+Dostie%2C+T.+A.+Richmond%2C+R.+A.+Arnaout%2C+...&amp;rft.date=2006-10-01&amp;rft.doi=10.1101%2Fgr.5571506&amp;rft.genre=journal&amp;rft.issn=1088-9051&amp;rft.issue=10&amp;rft.jtitle=Genome+Research&amp;rft.pages=1299-1309&amp;rft.pmc=1581439&amp;rft.pmid=16954542&amp;rft.volume=16" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-14"><span class="mw-cite-backlink"><a href="#cite_ref-14">↑</a></span> <span class="reference-text">Istvan Albert, Travis N. Mavrich, Lynn P. Tomsho, Ji Qi, Sara J. Zanton: <cite style="font-style:italic">Translational and rotational settings of H2A.Z nucleosomes across the Saccharomyces cerevisiae genome</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>446</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>7135</span>, März 2007, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>572–576</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/10.1038/nature05632">10.1038/nature05632</a></span>.<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Translational+and+rotational+settings+of+H2A.Z+nucleosomes+across+the+Saccharomyces+cerevisiae+genome&amp;rft.au=Istvan+Albert%2C+Travis+N.+Mavrich%2C+Lynn+P.+Tomsho%2C+...&amp;rft.date=2007-03&amp;rft.doi=10.1038%2Fnature05632&amp;rft.genre=journal&amp;rft.issue=7135&amp;rft.jtitle=Nature&amp;rft.pages=572-576&amp;rft.volume=446" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-15"><span class="mw-cite-backlink"><a href="#cite_ref-15">↑</a></span> <span class="reference-text">E. Lieberman-Aiden, N. L. van Berkum, L. Williams, M. Imakaev, T. Ragoczy: <cite style="font-style:italic">Comprehensive Mapping of Long-Range Interactions Reveals Folding Principles of the Human Genome</cite>. In: <cite style="font-style:italic"><a href="Science" title="Science">Science</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>326</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>5950</span>, 9.&nbsp;Oktober 2009, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>289–293</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/10.1126/science.1181369">10.1126/science.1181369</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/19815776?dopt=Abstract">PMID 19815776</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2858594/">PMC&nbsp;2858594</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Comprehensive+Mapping+of+Long-Range+Interactions+Reveals+Folding+Principles+of+the+Human+Genome&amp;rft.au=E.+Lieberman-Aiden%2C+N.+L.+van+Berkum%2C+L.+Williams%2C+...&amp;rft.date=2009-10-09&amp;rft.doi=10.1126%2Fscience.1181369&amp;rft.genre=journal&amp;rft.issue=5950&amp;rft.jtitle=Science&amp;rft.pages=289-293&amp;rft.pmc=2858594&amp;rft.pmid=19815776&amp;rft.volume=326" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-16"><span class="mw-cite-backlink"><a href="#cite_ref-16">↑</a></span> <span class="reference-text">Melissa J. Fullwood, Mei Hui Liu, You Fu Pan, Jun Liu, Han Xu: <cite style="font-style:italic">An oestrogen-receptor-α-bound human chromatin interactome</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>462</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>7269</span>, November 2009, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>58–64</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/10.1038/nature08497">10.1038/nature08497</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/19890323?dopt=Abstract">PMID 19890323</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2774924/">PMC&nbsp;2774924</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=An+oestrogen-receptor-%CE%B1-bound+human+chromatin+interactome&amp;rft.au=Melissa+J.+Fullwood%2C+Mei+Hui+Liu%2C+You+Fu+Pan%2C+...&amp;rft.date=2009-11&amp;rft.doi=10.1038%2Fnature08497&amp;rft.genre=journal&amp;rft.issue=7269&amp;rft.jtitle=Nature&amp;rft.pages=58-64&amp;rft.pmc=2774924&amp;rft.pmid=19890323&amp;rft.volume=462" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-17"><span class="mw-cite-backlink"><a href="#cite_ref-17">↑</a></span> <span class="reference-text">Jesse R. Dixon, Siddarth Selvaraj, Feng Yue, Audrey Kim, Yan Li: <cite style="font-style:italic">Topological domains in mammalian genomes identified by analysis of chromatin interactions</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>485</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>7398</span>, Mai 2012, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>376–380</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/10.1038/nature11082">10.1038/nature11082</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/22495300?dopt=Abstract">PMID 22495300</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3356448/">PMC&nbsp;3356448</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Topological+domains+in+mammalian+genomes+identified+by+analysis+of+chromatin+interactions&amp;rft.au=Jesse+R.+Dixon%2C+Siddarth+Selvaraj%2C+Feng+Yue%2C+...&amp;rft.date=2012-05&amp;rft.doi=10.1038%2Fnature11082&amp;rft.genre=journal&amp;rft.issue=7398&amp;rft.jtitle=Nature&amp;rft.pages=376-380&amp;rft.pmc=3356448&amp;rft.pmid=22495300&amp;rft.volume=485" style="display:none">&nbsp;</span></span>
</li>
<li id="cite_note-18"><span class="mw-cite-backlink"><a href="#cite_ref-18">↑</a></span> <span class="reference-text">Elphège P. Nora, Bryan R. Lajoie, Edda G. Schulz, Luca Giorgetti, Ikuhiro Okamoto: <cite style="font-style:italic">Spatial partitioning of the regulatory landscape of the X-inactivation centre</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em">&nbsp;</span>485</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em">&nbsp;</span>7398</span>, Mai 2012, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em">&nbsp;</span>381–385</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/10.1038/nature11049">10.1038/nature11049</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/22495304?dopt=Abstract">PMID 22495304</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3555144/">PMC&nbsp;3555144</a> (freier Volltext).<span class="Z3988" title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rfr_id=info:sid/de.wikipedia.org:Chromatin&amp;rft.atitle=Spatial+partitioning+of+the+regulatory+landscape+of+the+X-inactivation+centre&amp;rft.au=Elph%C3%A8ge+P.+Nora%2C+Bryan+R.+Lajoie%2C+Edda+G.+Schulz%2C+...&amp;rft.date=2012-05&amp;rft.doi=10.1038%2Fnature11049&amp;rft.genre=journal&amp;rft.issue=7398&amp;rft.jtitle=Nature&amp;rft.pages=381-385&amp;rft.pmc=3555144&amp;rft.pmid=22495304&amp;rft.volume=485" style="display:none">&nbsp;</span></span>
</li>
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