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<h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Protospacer Adjacent Motif</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"><p><b>Protospacer adjacent Motif</b> (PAM, ‚Vorabstandshalter-angrenzendes Motiv‘ bzw. Protospacer-Nachbarmotiv) ist in der Biochemie ein <a href="Sequenzmotiv" title="Sequenzmotiv">Sequenzmotiv</a> von <a href="DNA" class="mw-redirect" title="DNA">DNA</a>, an das die <a href="Endonuclease" class="mw-redirect" title="Endonuclease">Endonucleasen</a> vom Typ <a href="Cas9" title="Cas9">Cas</a> binden.<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>
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<div class="mw-heading mw-heading2"><h2 id="Eigenschaften">Eigenschaften</h2></div>
<p>Das PAM besteht aus 2 bis 6 <a href="Nukleotide" title="Nukleotide">Nukleotiden</a>. Das PAM ist die Bindungsstelle für Cas auf der zu schneidenden DNA. Cas9 kommt sowohl in der antiviralen Abwehr von Bakterien (<a href="CRISPR" title="CRISPR">CRISPR</a>), als auch im Zuge der <a href="CRISPR/Cas-Methode" title="CRISPR/Cas-Methode">CRISPR/Cas-Methode</a> zum <a href="Genome_Editing" title="Genome Editing">Genome Editing</a> vor. Ohne eine PAM hinter der zu schneidenden Sequenz erfolgt kein Schnitt der DNA.<sup id="cite_ref-pmid19246744_2-0" class="reference"><a href="#cite_note-pmid19246744-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid24578530_3-0" class="reference"><a href="#cite_note-pmid24578530-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid22745249_4-0" class="reference"><a href="#cite_note-pmid22745249-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid24476820_5-0" class="reference"><a href="#cite_note-pmid24476820-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In bakteriellen <a href="Genom" title="Genom">Genomen</a> kommt das PAM nicht in Kombination mit Erkennungssequenzen vor, weshalb es in Bakterien zur Erkennung fremder DNA dient.<sup id="cite_ref-pmid24076990_6-0" class="reference"><a href="#cite_note-pmid24076990-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Die von <a href="Cas9" title="Cas9">Cas9</a> aus <i><a href="Streptococcus_pyogenes" title="Streptococcus pyogenes">Streptococcus pyogenes</a></i> gebundene Sequenz ist 5'-NGG-3', mit N als beliebige <a href="Nukleinbase" class="mw-redirect" title="Nukleinbase">Nukleinbase</a> gefolgt von zwei <a href="Guanin" title="Guanin">Guaninen</a>.<sup id="cite_ref-pmid25079318_7-0" class="reference"><a href="#cite_note-pmid25079318-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Die Sequenz des PAM unterscheidet sich zwischen den Cas9-Varianten <i>Streptococcus pyogenes</i> und <i><a href="Neisseria_meningitidis" class="mw-redirect" title="Neisseria meningitidis">Neisseria meningitidis</a></i>, <i>Treponema denticola</i> und <i><a href="Streptococcus_thermophilus" title="Streptococcus thermophilus">Streptococcus thermophilus</a></i>.<sup id="cite_ref-pmid24076762_8-0" class="reference"><a href="#cite_note-pmid24076762-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Daneben kann auch ein Schnitt vor der Sequenz 5'-NGA-3' mit A als <a href="Adenosin" title="Adenosin">Adenosin</a> erfolgen.<sup id="cite_ref-pmid24956376_9-0" class="reference"><a href="#cite_note-pmid24956376-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Das PAM für <a href="Cas12b" title="Cas12b">Cas12b</a> aus <i><a href="Alicyclobacillus_acidoterrestris" title="Alicyclobacillus acidoterrestris">Alicyclobacillus acidoterrestris</a></i> besitzt die Sequenz 5'-TTN-3'.<sup id="cite_ref-uniprot-072_10-0" class="reference"><a href="#cite_note-uniprot-072-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>
</p><p>Ansätze des <a href="Protein-Engineering" title="Protein-Engineering">Protein-Engineering</a> zur Erweiterung der <a href="Substrat_(Biochemie)" title="Substrat (Biochemie)">Substratspezifität</a> von Cas9 durch Veränderung der PAM-Bindungsstelle wurden untersucht.<sup id="cite_ref-pmid26098369_11-0" class="reference"><a href="#cite_note-pmid26098369-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Das Cas9 von <i>Francisella novicida</i> wurde verändert, so dass die PAM 5'-YG-3' mit Y als ein beliebiges <a href="Pyrimidin" title="Pyrimidin">Pyrimidin</a> erkannt wird und ein Schnitt erfolgt.<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><sup id="cite_ref-pmid26875867_13-0" class="reference"><a href="#cite_note-pmid26875867-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Das <a href="Cpf1" title="Cpf1">Cas12a</a> (vormals Cpf1<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>) von <i>Francisella novicida</i> bindet die PAM 5'-TTTN-3' mit T als <a href="Thymidin" class="mw-redirect" title="Thymidin">Thymidin</a><sup id="cite_ref-pmid26422227_15-0" class="reference"><a href="#cite_note-pmid26422227-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> oder 5'-YTN-3'.<sup id="cite_ref-pmid27096362_16-0" class="reference"><a href="#cite_note-pmid27096362-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> <a href="Cas13" title="Cas13">Cas13</a> Proteine, wie Cas13a von <i>Leptotrichia shahii (vormals C2c2),</i> binden und schneiden <a href="Ribonukleins%C3%A4ure" title="Ribonukleinsäure">RNA</a> anstatt <a href="Desoxyribonukleins%C3%A4ure" title="Desoxyribonukleinsäure">DNA</a> und binden zum Teil an eine <i>Protospacer Flanking Site</i> (PFS) anstatt an ein PAM. Diese PFS besteht bei Cas13a von <i>Leptotrichia shahii</i> zum Beispiel aus einem beliebigen <a href="Nukleotide" title="Nukleotide">Nukleotid</a> außer <a href="Guanosin" title="Guanosin">Guanosin</a>.<sup id="cite_ref-pmid27256883_17-0" class="reference"><a href="#cite_note-pmid27256883-17"><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> Es wurde jedoch gezeigt, dass Cas13 Proteine anderer Spezies keine PFS zur Bindung benötigen.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup>
</p>
<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">S. A. Shah, S. Erdmann, F. J. Mojica, R. A. Garrett: <i>Protospacer recognition motifs: mixed identities and functional diversity.</i> In: <i>RNA biology.</i> Band 10, Nummer 5, Mai 2013, S. 891–899, <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/10.4161/rna.23764">10.4161/rna.23764</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/23403393?dopt=Abstract">PMID 23403393</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3737346/">PMC 3737346</a> (freier Volltext).</span>
</li>
<li id="cite_note-pmid19246744-2"><span class="mw-cite-backlink"><a href="#cite_ref-pmid19246744_2-0">↑</a></span> <span class="reference-text">Mojica FJ, Díez-Villaseñor C, García-Martínez J, Almendros C: <cite style="font-style:italic">Short motif sequences determine the targets of the prokaryotic CRISPR defence system</cite>. In: <cite style="font-style:italic"><a href="Microbiology" title="Microbiology">Microbiology</a></cite>. 155. Jahrgang, Pt 3, 2009, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>733–740</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/10.1099/mic.0.023960-0">10.1099/mic.0.023960-0</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/19246744?dopt=Abstract">PMID 19246744</a> (<a rel="nofollow" class="external text" href="http://mic.sgmjournals.org/content/155/3/733.long">sgmjournals.org</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:Protospacer+Adjacent+Motif&rft.atitle=Short+motif+sequences+determine+the+targets+of+the+prokaryotic+CRISPR+defence+system&rft.au=Mojica+FJ%2C+D%C3%ADez-Villase%C3%B1or+C%2C+Garc%C3%ADa-Mart%C3%ADnez+J%2C+...&rft.date=2009&rft.doi=10.1099%2Fmic.0.023960-0&rft.genre=journal&rft.issue=Pt+3&rft.jtitle=Microbiology&rft.pages=733-740&rft.pmid=19246744&rft.volume=155.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid24578530-3"><span class="mw-cite-backlink"><a href="#cite_ref-pmid24578530_3-0">↑</a></span> <span class="reference-text">Shah SA, Erdmann S, Mojica FJ, Garrett RA: <cite class="lang" lang="en" dir="auto" style="font-style:italic">Protospacer recognition motifs: mixed identities and functional diversity</cite>. In: <cite class="lang" lang="en" dir="auto" style="font-style:italic"><a href="RNA_Biology" title="RNA Biology">RNA Biology</a></cite>. 10. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>5</span>, 2013, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>891–899</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/10.4161/rna.23764">10.4161/rna.23764</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/23403393?dopt=Abstract">PMID 23403393</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3737346/">PMC 3737346</a> (freier Volltext) – (englisch, <a rel="nofollow" class="external text" href="https://web.archive.org/web/20140904034408/https://www.landesbioscience.com/journals/rnabiology/article/23764/">landesbioscience.com</a> (<span class="webarchiv-memento"><a href="Webarchivierung#Begrifflichkeiten" title="Webarchivierung">Memento</a></span> des <style data-mw-deduplicate="TemplateStyles:r250917974">
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<li id="cite_note-pmid22745249-4"><span class="mw-cite-backlink"><a href="#cite_ref-pmid22745249_4-0">↑</a></span> <span class="reference-text">Jinek M, Chylinski K, Fonfara I, Hauer M, Doudna JA, Charpentier E: <cite style="font-style:italic">A programmable dual-RNA-guided DNA endonuclease in adaptive bacterial immunity</cite>. In: <cite style="font-style:italic"><a href="Science" title="Science">Science</a></cite>. 337. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>6096</span>, 2012, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>816–821</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.1225829">10.1126/science.1225829</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/22745249?dopt=Abstract">PMID 22745249</a> (<a rel="nofollow" class="external text" href="http://www.sciencemag.org/content/337/6096/816.long">sciencemag.org</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:Protospacer+Adjacent+Motif&rft.atitle=A+programmable+dual-RNA-guided+DNA+endonuclease+in+adaptive+bacterial+immunity&rft.au=Jinek+M%2C+Chylinski+K%2C+Fonfara+I%2C+...&rft.date=2012&rft.doi=10.1126%2Fscience.1225829&rft.genre=journal&rft.issue=6096&rft.jtitle=Science&rft.pages=816-821&rft.pmid=22745249&rft.volume=337.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid24476820-5"><span class="mw-cite-backlink"><a href="#cite_ref-pmid24476820_5-0">↑</a></span> <span class="reference-text">Sternberg SH, Redding S, Jinek M, Greene EC, Doudna JA: <cite style="font-style:italic">DNA interrogation by the CRISPR RNA-guided endonuclease Cas9</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. 507. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>7490</span>, 2014, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>62–67</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/nature13011">10.1038/nature13011</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24476820?dopt=Abstract">PMID 24476820</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4106473/">PMC 4106473</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:Protospacer+Adjacent+Motif&rft.atitle=DNA+interrogation+by+the+CRISPR+RNA-guided+endonuclease+Cas9&rft.au=Sternberg+SH%2C+Redding+S%2C+Jinek+M%2C+...&rft.date=2014&rft.doi=10.1038%2Fnature13011&rft.genre=journal&rft.issue=7490&rft.jtitle=Nature&rft.pages=62-67&rft.pmc=4106473&rft.pmid=24476820&rft.volume=507.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid24076990-6"><span class="mw-cite-backlink"><a href="#cite_ref-pmid24076990_6-0">↑</a></span> <span class="reference-text">Mali P, Esvelt KM, Church GM: <cite style="font-style:italic">Cas9 as a versatile tool for engineering biology</cite>. In: <cite style="font-style:italic"><a href="Nature_Methods" title="Nature Methods">Nature Methods</a></cite>. 10. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>10</span>, 2013, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>957–963</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/nmeth.2649">10.1038/nmeth.2649</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24076990?dopt=Abstract">PMID 24076990</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4051438/">PMC 4051438</a> (freier Volltext) – (<a rel="nofollow" class="external text" href="http://www.nature.com/nmeth/journal/v10/n10/full/nmeth.2649.html">nature.com</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:Protospacer+Adjacent+Motif&rft.atitle=Cas9+as+a+versatile+tool+for+engineering+biology&rft.au=Mali+P%2C+Esvelt+KM%2C+Church+GM&rft.date=2013&rft.doi=10.1038%2Fnmeth.2649&rft.genre=journal&rft.issue=10&rft.jtitle=Nature+Methods&rft.pages=957-963&rft.pmc=4051438&rft.pmid=24076990&rft.volume=10.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid25079318-7"><span class="mw-cite-backlink"><a href="#cite_ref-pmid25079318_7-0">↑</a></span> <span class="reference-text">Anders C, Niewoehner O, Duerst A, Jinek M: <cite style="font-style:italic">Structural basis of PAM-dependent target DNA recognition by the Cas9 endonuclease</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. 513. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>7519</span>, 2014, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>569–573</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/nature13579">10.1038/nature13579</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/25079318?dopt=Abstract">PMID 25079318</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4176945/">PMC 4176945</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:Protospacer+Adjacent+Motif&rft.atitle=Structural+basis+of+PAM-dependent+target+DNA+recognition+by+the+Cas9+endonuclease&rft.au=Anders+C%2C+Niewoehner+O%2C+Duerst+A%2C+...&rft.date=2014&rft.doi=10.1038%2Fnature13579&rft.genre=journal&rft.issue=7519&rft.jtitle=Nature&rft.pages=569-573&rft.pmc=4176945&rft.pmid=25079318&rft.volume=513.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid24076762-8"><span class="mw-cite-backlink"><a href="#cite_ref-pmid24076762_8-0">↑</a></span> <span class="reference-text">Esvelt KM, Mali P, Braff JL, Moosburner M, Yaung SJ, Church GM: <cite style="font-style:italic">Orthogonal Cas9 proteins for RNA-guided gene regulation and editing</cite>. In: <cite style="font-style:italic"><a href="Nature_Methods" title="Nature Methods">Nature Methods</a></cite>. 10. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>11</span>, 2013, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>1116–1123</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/nmeth.2681">10.1038/nmeth.2681</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24076762?dopt=Abstract">PMID 24076762</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3844869/">PMC 3844869</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:Protospacer+Adjacent+Motif&rft.atitle=Orthogonal+Cas9+proteins+for+RNA-guided+gene+regulation+and+editing&rft.au=Esvelt+KM%2C+Mali+P%2C+Braff+JL%2C+...&rft.date=2013&rft.doi=10.1038%2Fnmeth.2681&rft.genre=journal&rft.issue=11&rft.jtitle=Nature+Methods&rft.pages=1116-1123&rft.pmc=3844869&rft.pmid=24076762&rft.volume=10.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid24956376-9"><span class="mw-cite-backlink"><a href="#cite_ref-pmid24956376_9-0">↑</a></span> <span class="reference-text">Zhang Y, Ge X, Yang F, Zhang L, Zheng J, Tan X, Jin ZB, Qu J, Gu F: <cite style="font-style:italic">Comparison of non-canonical PAMs for CRISPR/Cas9-mediated DNA cleavage in human cells</cite>. In: <cite style="font-style:italic"><a href="Scientific_Reports" title="Scientific Reports">Scientific Reports</a></cite>. 4. Jahrgang, 2014, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>5405</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/srep05405">10.1038/srep05405</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/24956376?dopt=Abstract">PMID 24956376</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4066725/">PMC 4066725</a> (freier Volltext).<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:Protospacer+Adjacent+Motif&rft.atitle=Comparison+of+non-canonical+PAMs+for+CRISPR%2FCas9-mediated+DNA+cleavage+in+human+cells&rft.au=Zhang+Y%2C+Ge+X%2C+Yang+F%2C+...&rft.btitle=Scientific+Reports&rft.date=2014&rft.doi=10.1038%2Fsrep05405&rft.genre=book&rft.pages=5405&rft.pmc=4066725&rft.pmid=24956376&rft.volume=4.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-uniprot-072-10"><span class="mw-cite-backlink"><a href="#cite_ref-uniprot-072_10-0">↑</a></span> <span class="reference-text"><a href="UniProt" title="UniProt">UniProt</a> <a rel="nofollow" class="external text" href="https://www.uniprot.org/uniprotkb/T0D7A2">cas12b - CRISPR-associated endonuclease Cas12b - Alicyclobacillus acidoterrestris (strain ATCC 49025 / DSM 3922 / CIP 106132 / NCIMB 13137 / GD3B) - cas12b gene</a>. Abgerufen am 24. Januar 2019.</span>
</li>
<li id="cite_note-pmid26098369-11"><span class="mw-cite-backlink"><a href="#cite_ref-pmid26098369_11-0">↑</a></span> <span class="reference-text">Kleinstiver BP, Prew MS, Tsai SQ, Topkar VV, Nguyen NT, Zheng Z, Gonzales AP, Li Z, Peterson RT, Yeh JR, Aryee MJ, Joung JK: <cite style="font-style:italic">Engineered CRISPR-Cas9 nucleases with altered PAM specificities</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. 523. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>7561</span>, 2015, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>481–485</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/nature14592">10.1038/nature14592</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/26098369?dopt=Abstract">PMID 26098369</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4540238/">PMC 4540238</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:Protospacer+Adjacent+Motif&rft.atitle=Engineered+CRISPR-Cas9+nucleases+with+altered+PAM+specificities&rft.au=Kleinstiver+BP%2C+Prew+MS%2C+Tsai+SQ%2C+...&rft.date=2015&rft.doi=10.1038%2Fnature14592&rft.genre=journal&rft.issue=7561&rft.jtitle=Nature&rft.pages=481-485&rft.pmc=4540238&rft.pmid=26098369&rft.volume=523.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-12"><span class="mw-cite-backlink"><a href="#cite_ref-12">↑</a></span> <span class="reference-text"><span class="cite"><a rel="nofollow" class="external text" href="http://www.genome.jp/kegg/catalog/codes1.html"><i>Nucleotide Codes, Amino Acid Codes, and Genetic Codes.</i></a> KEGG: Kyoto Encyclopedia of Genes and Genomes, 15. Juli 2014,<span class="Abrufdatum"> abgerufen am 6. April 2016</span>.</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%3AProtospacer+Adjacent+Motif&rft.title=Nucleotide+Codes%2C+Amino+Acid+Codes%2C+and+Genetic+Codes&rft.description=Nucleotide+Codes%2C+Amino+Acid+Codes%2C+and+Genetic+Codes&rft.identifier=&rft.publisher=KEGG%3A+Kyoto+Encyclopedia+of+Genes+and+Genomes&rft.date=2014-07-15"> </span></span>
</li>
<li id="cite_note-pmid26875867-13"><span class="mw-cite-backlink"><a href="#cite_ref-pmid26875867_13-0">↑</a></span> <span class="reference-text">Hirano H, Gootenberg JS, Horii T, Abudayyeh OO, Kimura M, Hsu PD, Nakane T, Ishitani R, Hatada I, Zhang F, Nishimasu H, Nureki O: <cite style="font-style:italic">Structure and Engineering of Francisella novicida Cas9</cite>. In: <cite style="font-style:italic"><a href="Cell_(Zeitschrift)" title="Cell (Zeitschrift)">Cell</a></cite>. 164. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>5</span>, 2016, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>950–961</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.cell.2016.01.039">10.1016/j.cell.2016.01.039</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/26875867?dopt=Abstract">PMID 26875867</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4899972/">PMC 4899972</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:Protospacer+Adjacent+Motif&rft.atitle=Structure+and+Engineering+of+Francisella+novicida+Cas9&rft.au=Hirano+H%2C+Gootenberg+JS%2C+Horii+T%2C+...&rft.date=2016&rft.doi=10.1016%2Fj.cell.2016.01.039&rft.genre=journal&rft.issue=5&rft.jtitle=Cell&rft.pages=950-961&rft.pmc=4899972&rft.pmid=26875867&rft.volume=164.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-14"><span class="mw-cite-backlink"><a href="#cite_ref-14">↑</a></span> <span class="reference-text">Kira S. Makarova, Yuri I. Wolf, Eugene V. Koonin: <cite style="font-style:italic">Evolutionary Classification of CRISPR‐Cas Systems</cite>. In: <cite style="font-style:italic">Crispr</cite>. 1. Auflage. Wiley, 2022, ISBN 978-1-68367-037-7, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>13–38</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/10.1002/9781683673798.ch2">10.1002/9781683673798.ch2</a></span> (<a rel="nofollow" class="external text" href="https://onlinelibrary.wiley.com/doi/10.1002/9781683673798.ch2">wiley.com</a> [abgerufen am 25. August 2024]).<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:Protospacer+Adjacent+Motif&rft.atitle=Evolutionary+Classification+of+CRISPR%E2%80%90Cas+Systems&rft.au=Kira+S.+Makarova%2C+Yuri+I.+Wolf%2C+Eugene+V.+Koonin&rft.btitle=Crispr&rft.date=2022-08-30&rft.doi=10.1002%2F9781683673798.ch2&rft.edition=1&rft.genre=book&rft.isbn=9781683670377&rft.pages=13-38&rft.pub=Wiley" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid26422227-15"><span class="mw-cite-backlink"><a href="#cite_ref-pmid26422227_15-0">↑</a></span> <span class="reference-text">Zetsche B, Gootenberg JS, Abudayyeh OO, Slaymaker IM, Makarova KS, Essletzbichler P, Volz SE, Joung J, van der Oost J, Regev A, Koonin EV, Zhang F: <cite style="font-style:italic">Cpf1 is a single RNA-guided endonuclease of a class 2 CRISPR-Cas system</cite>. In: <cite style="font-style:italic"><a href="Cell_(Zeitschrift)" title="Cell (Zeitschrift)">Cell</a></cite>. 163. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>3</span>, 2015, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>759–771</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.cell.2015.09.038">10.1016/j.cell.2015.09.038</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/26422227?dopt=Abstract">PMID 26422227</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:Protospacer+Adjacent+Motif&rft.atitle=Cpf1+is+a+single+RNA-guided+endonuclease+of+a+class+2+CRISPR-Cas+system&rft.au=Zetsche+B%2C+Gootenberg+JS%2C+Abudayyeh+OO%2C+...&rft.date=2015&rft.doi=10.1016%2Fj.cell.2015.09.038&rft.genre=journal&rft.issue=3&rft.jtitle=Cell&rft.pages=759-771&rft.pmid=26422227&rft.volume=163.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid27096362-16"><span class="mw-cite-backlink"><a href="#cite_ref-pmid27096362_16-0">↑</a></span> <span class="reference-text">Fonfara I, Richter H, Bratovič M, Le Rhun A, Charpentier E: <cite style="font-style:italic">The CRISPR-associated DNA-cleaving enzyme Cpf1 also processes precursor CRISPR RNA</cite>. In: <cite style="font-style:italic"><a href="Nature" title="Nature">Nature</a></cite>. 532. Jahrgang, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>7600</span>, 2016, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>517–521</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/nature17945">10.1038/nature17945</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/27096362?dopt=Abstract">PMID 27096362</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:Protospacer+Adjacent+Motif&rft.atitle=The+CRISPR-associated+DNA-cleaving+enzyme+Cpf1+also+processes+precursor+CRISPR+RNA&rft.au=Fonfara+I%2C+Richter+H%2C+Bratovi%C4%8D+M%2C+...&rft.date=2016&rft.doi=10.1038%2Fnature17945&rft.genre=journal&rft.issue=7600&rft.jtitle=Nature&rft.pages=517-521&rft.pmid=27096362&rft.volume=532.+Jahrgang" style="display:none"> </span></span>
</li>
<li id="cite_note-pmid27256883-17"><span class="mw-cite-backlink"><a href="#cite_ref-pmid27256883_17-0">↑</a></span> <span class="reference-text">Abudayyeh OO, Gootenberg JS, Konermann S, Joung J, Slaymaker IM, Cox DB, Shmakov S, Makarova KS, Semenova E, Minakhin L, Severinov K, Regev A, Lander ES, Koonin EV, Zhang F: <cite style="font-style:italic">C2c2 is a single-component programmable RNA-guided RNA-targeting CRISPR effector</cite>. In: <cite style="font-style:italic"><a href="Science" title="Science">Science</a></cite>. 2016, <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.aaf5573">10.1126/science.aaf5573</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/27256883?dopt=Abstract">PMID 27256883</a>.<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:Protospacer+Adjacent+Motif&rft.atitle=C2c2+is+a+single-component+programmable+RNA-guided+RNA-targeting+CRISPR+effector&rft.au=Abudayyeh+OO%2C+Gootenberg+JS%2C+Konermann+S%2C+...&rft.btitle=Science&rft.date=2016&rft.doi=10.1126%2Fscience.aaf5573&rft.genre=book&rft.pmid=27256883" style="display:none"> </span></span>
</li>
<li id="cite_note-18"><span class="mw-cite-backlink"><a href="#cite_ref-18">↑</a></span> <span class="reference-text">Kira S. Makarova, Feng Zhang, Eugene V. Koonin: <cite style="font-style:italic">SnapShot: Class 2 CRISPR-Cas Systems</cite>. In: <cite style="font-style:italic">Cell</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em"> </span>168</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>1–2</span>, Januar 2017, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>328–328.e1</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.cell.2016.12.038">10.1016/j.cell.2016.12.038</a></span> (<a rel="nofollow" class="external text" href="https://linkinghub.elsevier.com/retrieve/pii/S0092867416317548">elsevier.com</a> [abgerufen am 17. Mai 2020]).<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:Protospacer+Adjacent+Motif&rft.atitle=SnapShot%3A+Class+2+CRISPR-Cas+Systems&rft.au=Kira+S.+Makarova%2C+Feng+Zhang%2C+Eugene+V.+Koonin&rft.date=2017-01&rft.doi=10.1016%2Fj.cell.2016.12.038&rft.genre=journal&rft.issue=1-2&rft.jtitle=Cell&rft.pages=328-328.e1&rft.volume=168" style="display:none"> </span></span>
</li>
<li id="cite_note-19"><span class="mw-cite-backlink"><a href="#cite_ref-19">↑</a></span> <span class="reference-text">Omar O. Abudayyeh, Jonathan S. Gootenberg, Patrick Essletzbichler, Shuo Han, Julia Joung: <cite style="font-style:italic">RNA targeting with CRISPR–Cas13</cite>. In: <cite style="font-style:italic">Nature</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em"> </span>550</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>7675</span>, Oktober 2017, <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%220028-0836%22&key=cql">0028-0836</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>280–284</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/nature24049">10.1038/nature24049</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/28976959?dopt=Abstract">PMID 28976959</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5706658/">PMC 5706658</a> (freier Volltext) – (<a rel="nofollow" class="external text" href="http://www.nature.com/articles/nature24049">nature.com</a> [abgerufen am 17. Mai 2020]).<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:Protospacer+Adjacent+Motif&rft.atitle=RNA+targeting+with+CRISPR-Cas13&rft.au=Omar+O.+Abudayyeh%2C+Jonathan+S.+Gootenberg%2C+Patrick+Essletzbichler%2C+...&rft.date=2017-10&rft.doi=10.1038%2Fnature24049&rft.genre=journal&rft.issn=0028-0836&rft.issue=7675&rft.jtitle=Nature&rft.pages=280-284&rft.pmc=5706658&rft.pmid=28976959&rft.volume=550" style="display:none"> </span></span>
</li>
<li id="cite_note-20"><span class="mw-cite-backlink"><a href="#cite_ref-20">↑</a></span> <span class="reference-text">David B. T. Cox, Jonathan S. Gootenberg, Omar O. Abudayyeh, Brian Franklin, Max J. Kellner: <cite style="font-style:italic">RNA editing with CRISPR-Cas13</cite>. In: <cite style="font-style:italic">Science</cite>. <span style="white-space:nowrap">Band<span style="display:inline-block;width:.2em"> </span>358</span>, <span style="white-space:nowrap">Nr.<span style="display:inline-block;width:.2em"> </span>6366</span>, 24. November 2017, <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%220036-8075%22&key=cql">0036-8075</a></span>, <span style="white-space:nowrap">S.<span style="display:inline-block;width:.2em"> </span>1019–1027</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.aaq0180">10.1126/science.aaq0180</a></span>, <a class="external mw-magiclink-pmid" rel="nofollow" href="https://www.ncbi.nlm.nih.gov/pubmed/29070703?dopt=Abstract">PMID 29070703</a>, <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5793859/">PMC 5793859</a> (freier Volltext) – (<a rel="nofollow" class="external text" href="https://www.sciencemag.org/lookup/doi/10.1126/science.aaq0180">sciencemag.org</a> [abgerufen am 17. Mai 2020]).<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:Protospacer+Adjacent+Motif&rft.atitle=RNA+editing+with+CRISPR-Cas13&rft.au=David+B.+T.+Cox%2C+Jonathan+S.+Gootenberg%2C+Omar+O.+Abudayyeh%2C+...&rft.date=2017-11-24&rft.doi=10.1126%2Fscience.aaq0180&rft.genre=journal&rft.issn=0036-8075&rft.issue=6366&rft.jtitle=Science&rft.pages=1019-1027&rft.pmc=5793859&rft.pmid=29070703&rft.volume=358" style="display:none"> </span></span>
</li>
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