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class="vector-toc-numb">2</span> <span>Gene knockout by mutation</span> </div> </a> <ul id="toc-Gene_knockout_by_mutation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gene_silencing" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Gene_silencing"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Gene silencing</span> </div> </a> <button aria-controls="toc-Gene_silencing-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Gene silencing subsection</span> </button> <ul id="toc-Gene_silencing-sublist" class="vector-toc-list"> <li id="toc-Homologous_recombination" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Homologous_recombination"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Homologous recombination</span> </div> </a> <ul id="toc-Homologous_recombination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Site-specific_nucleases" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Site-specific_nucleases"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Site-specific nucleases</span> </div> </a> <ul id="toc-Site-specific_nucleases-sublist" class="vector-toc-list"> <li id="toc-Zinc-fingers" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Zinc-fingers"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.1</span> <span>Zinc-fingers</span> </div> </a> <ul id="toc-Zinc-fingers-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-TALENS" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#TALENS"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.2</span> <span>TALENS</span> </div> </a> <ul id="toc-TALENS-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-CRISPR/Cas9" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#CRISPR/Cas9"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2.3</span> <span>CRISPR/Cas9</span> </div> </a> <ul id="toc-CRISPR/Cas9-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Knock-in" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Knock-in"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Knock-in</span> </div> </a> <ul id="toc-Knock-in-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Types" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Types"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Types</span> </div> </a> <button aria-controls="toc-Types-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Types subsection</span> </button> <ul id="toc-Types-sublist" class="vector-toc-list"> <li id="toc-Conditional_knockouts" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Conditional_knockouts"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Conditional knockouts</span> </div> </a> <ul id="toc-Conditional_knockouts-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Use" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Use"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Use</span> </div> </a> <ul id="toc-Use-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> 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class="firstHeading mw-first-heading"><span class="mw-page-title-main">Gene knockout</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 22 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-22" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">22 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D8%B9%D8%B7%D9%8A%D9%84_%D9%85%D9%88%D8%B1%D8%AB%D8%A9" title="تعطيل مورثة – Arabic" lang="ar" hreflang="ar" data-title="تعطيل مورثة" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9D%D0%BE%D0%BA%D0%B0%D1%83%D1%82_(%D0%B3%D0%B5%D0%BD%D0%B5%D1%82%D0%B8%D0%BA%D0%B0)" title="Нокаут (генетика) – Bulgarian" lang="bg" hreflang="bg" data-title="Нокаут (генетика)" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Genski_nokaut" title="Genski nokaut – Bosnian" lang="bs" hreflang="bs" data-title="Genski nokaut" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Organisme_genoanul%C2%B7lat" title="Organisme genoanul·lat – Catalan" lang="ca" hreflang="ca" data-title="Organisme genoanul·lat" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Genov%C3%BD_knockout" title="Genový knockout – Czech" lang="cs" hreflang="cs" data-title="Genový knockout" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Gen-Knockout" title="Gen-Knockout – German" lang="de" hreflang="de" data-title="Gen-Knockout" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%93%CE%BF%CE%BD%CE%B9%CE%B4%CE%B9%CE%B1%CE%BA%CE%AE_%CE%B1%CF%80%CE%B5%CE%BD%CE%B5%CF%81%CE%B3%CE%BF%CF%80%CE%BF%CE%AF%CE%B7%CF%83%CE%B7" title="Γονιδιακή απενεργοποίηση – Greek" lang="el" hreflang="el" data-title="Γονιδιακή απενεργοποίηση" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Bloqueo_de_genes" title="Bloqueo de genes – Spanish" lang="es" hreflang="es" data-title="Bloqueo de genes" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%B3%D8%B1%DA%A9%D9%88%D8%A8_%DA%98%D9%86" title="سرکوب ژن – Persian" lang="fa" hreflang="fa" data-title="سرکوب ژن" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Knock-out_(g%C3%A9n%C3%A9tique)" title="Knock-out (génétique) – French" lang="fr" hreflang="fr" data-title="Knock-out (génétique)" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Knockout_de_xenes" title="Knockout de xenes – Galician" lang="gl" hreflang="gl" data-title="Knockout de xenes" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%9C%A0%EC%A0%84%EC%9E%90_%EC%A0%9C%EA%B1%B0" title="유전자 제거 – Korean" lang="ko" hreflang="ko" data-title="유전자 제거" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B3%D5%A5%D5%B6%D5%AB_%D5%B6%D5%B8%D5%AF%D5%A1%D5%B8%D6%82%D5%BF" title="Գենի նոկաուտ – Armenian" lang="hy" hreflang="hy" data-title="Գենի նոկաուտ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Gensvipting" title="Gensvipting – Icelandic" lang="is" hreflang="is" data-title="Gensvipting" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target"><span>Íslenska</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%A0%D7%95%D7%A7%D7%90%D7%90%D7%95%D7%98_%D7%92%D7%A0%D7%98%D7%99" title="נוקאאוט גנטי – Hebrew" lang="he" hreflang="he" data-title="נוקאאוט גנטי" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Knock-out_(biologie)" title="Knock-out (biologie) – Dutch" lang="nl" hreflang="nl" data-title="Knock-out (biologie)" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E9%81%BA%E4%BC%9D%E5%AD%90%E3%83%8E%E3%83%83%E3%82%AF%E3%82%A2%E3%82%A6%E3%83%88" title="遺伝子ノックアウト – Japanese" lang="ja" hreflang="ja" data-title="遺伝子ノックアウト" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Nocaute_de_genes" title="Nocaute de genes – Portuguese" lang="pt" hreflang="pt" data-title="Nocaute de genes" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9D%D0%BE%D0%BA%D0%B0%D1%83%D1%82_%D0%B3%D0%B5%D0%BD%D0%B0" title="Нокаут гена – Russian" lang="ru" hreflang="ru" data-title="Нокаут гена" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Gene_knockout" title="Gene knockout – Simple English" lang="en-simple" hreflang="en-simple" data-title="Gene knockout" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a 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div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">"Triple knockout" redirects here. For the playoff tournament, see <a href="/wiki/Tournament#Knockout_tournaments" title="Tournament">Tournament § Knockout tournaments</a>.</div> <p><b>Gene knockouts</b> (also known as <b>gene deletion</b> or <b>gene inactivation</b>) are a widely used genetic engineering technique that involves the <a href="/wiki/Gene_targeting" title="Gene targeting">targeted</a> removal or inactivation of a specific gene within an organism's genome. This can be done through a variety of methods, including <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombination</a>, <a href="/wiki/CRISPR_gene_editing" title="CRISPR gene editing">CRISPR-Cas9</a>, and <a href="/wiki/Transcription_activator-like_effector_nuclease" title="Transcription activator-like effector nuclease">TALENs</a>. </p><p>One of the main advantages of gene knockouts is that they allow researchers to study the function of a specific gene in vivo, and to understand the role of the gene in normal development and physiology as well as in the pathology of diseases. By studying the <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> of the organism with the knocked out gene, researchers can gain insights into the biological processes that the gene is involved in. </p><p>There are two main types of gene knockouts: complete and conditional. A complete gene knockout permanently inactivates the gene, while a conditional gene knockout allows for the gene to be turned off and on at specific times or in specific tissues. Conditional knockouts are particularly useful for studying developmental processes and for understanding the role of a gene in specific cell types or tissues. </p><p>Gene knockouts have been widely used in many different organisms, including bacteria, yeast, fruit flies, zebrafish, and mice. In mice, gene knockouts are commonly used to study the function of specific genes in development, physiology, and cancer research. </p><p>The use of gene knockouts in mouse models has been particularly valuable in the study of human diseases. For example, gene knockouts in mice have been used to study the role of specific genes in cancer, neurological disorders, immune disorders, and metabolic disorders. </p><p>However, gene knockouts also have some limitations. For example, the loss of a single gene may not fully mimic the effects of a genetic disorder, and the knockouts may have unintended effects on other genes or pathways. Additionally, gene knockouts are not always a good model for human disease as the mouse genome is not identical to the human genome, and mouse physiology is different from human physiology. </p><p>The KO technique is essentially the opposite of a <a href="/wiki/Gene_knock-in" title="Gene knock-in">gene knock-in</a>. Knocking out two genes simultaneously in an organism is known as a <b>double knockout</b> (<b>DKO</b>). Similarly the terms <b>triple knockout</b> (<b>TKO</b>) and <b>quadruple knockouts</b> (<b>QKO</b>) are used to describe three or four knocked out genes, respectively. However, one needs to distinguish between <a href="/wiki/Zygosity" title="Zygosity">heterozygous and homozygous</a> KOs. In the former, only one of two gene copies (<a href="/wiki/Allele" title="Allele">alleles</a>) is knocked out, in the latter both are knocked out. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Methods">Methods</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=1" title="Edit section: Methods"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div><p> Knockouts are accomplished through a variety of techniques. Originally, <b>naturally occurring <a href="/wiki/Mutation" title="Mutation">mutations</a></b> were identified and then gene loss or inactivation had to be established by <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> or other methods.<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><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Knockout_Mice5006-300.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e1/Knockout_Mice5006-300.jpg/220px-Knockout_Mice5006-300.jpg" decoding="async" width="220" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e1/Knockout_Mice5006-300.jpg/330px-Knockout_Mice5006-300.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e1/Knockout_Mice5006-300.jpg/440px-Knockout_Mice5006-300.jpg 2x" data-file-width="3008" data-file-height="1960" /></a><figcaption>A laboratory mouse in which a gene affecting hair growth has been knocked out (left), is shown next to a normal lab mouse.</figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Gene_knockout_by_mutation">Gene knockout by mutation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=2" title="Edit section: Gene knockout by mutation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Gene knockout by mutation is commonly carried out in bacteria. An early instance of the use of this technique in <i>Escherichia coli</i> was published in 1989 by Hamilton, et al.<sup id="cite_ref-Hamilton_2-0" class="reference"><a href="#cite_note-Hamilton-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In this experiment, two sequential recombinations were used to delete the gene. This work established the feasibility of removing or replacing a functional gene in bacteria. That method has since been developed for other organisms, particularly research animals, like mice. Knockout mice are commonly used to study genes with human equivalents that may have significance for disease. An example of a study using knockout mice is an investigation of the roles of <a href="/wiki/XIRP2" title="XIRP2">Xirp</a> proteins in Sudden Unexplained Nocturnal Death Syndrome (SUNDS) and Brugada Syndrome in the Chinese Han Population.<sup id="cite_ref-Huang_3-0" class="reference"><a href="#cite_note-Huang-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Gene_silencing">Gene silencing</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=3" title="Edit section: Gene silencing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>For gene knockout investigations, <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a> (RNAi), a more recent method, also known as gene silencing, has gained popularity. In RNA interference (RNAi), messenger RNA for a particular gene is inactivated using small interfering RNA (siRNA) or short hairpin RNA (shRNA). This effectively stops the gene from being expressed. Oncogenes like Bcl-2 and p53, as well as genes linked to neurological disease, genetic disorders, and viral infections, have all been targeted for gene silencing utilizing RNA interference (RNAi).<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> </p> <div class="mw-heading mw-heading3"><h3 id="Homologous_recombination">Homologous recombination</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=4" title="Edit section: Homologous recombination"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Homologous_recombination" title="Homologous recombination">Homologous recombination</a></div> <p>Homologous recombination is the exchange of genes between two DNA strands that include extensive regions of base sequences that are identical to one another. In eukaryotic species, bacteria, and some viruses, homologous recombination happens spontaneously and is a useful tool in genetic engineering. Homologous recombination, which takes place during meiosis in eukaryotes, is essential for the repair of double-stranded DNA breaks and promotes genetic variation by allowing the movement of genetic information during chromosomal crossing. Homologous recombination, a key DNA repair mechanism in bacteria, enables the insertion of genetic material acquired through horizontal transfer of genes and transformation into DNA. Homologous recombination in viruses influences the course of viral evolution. Homologous recombination, a type of gene targeting used in genetic engineering, involves the introduction of an engineered mutation into a particular gene in order to learn more about the function of that gene. This method involves inserting foreign DNA into a cell that has a sequence similar to the target gene while being flanked by sequences that are the same upstream and downstream of the target gene. The target gene's DNA is substituted with the foreign DNA sequence during replication when the cell detects the similar flanking regions as homologues. The target gene is "knocked out" by the exchange. By using this technique to target particular alleles in embryonic stem cells in mice, it is possible to create knockout mice. With the aid of gene targeting, numerous mouse genes have been shut down, leading to the creation of hundreds of distinct mouse models of various human diseases, such as cancer, diabetes, cardiovascular diseases, and neurological disorders.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (December 2023)">citation needed</span></a></i>]</sup> Mario Capecchi, Sir Martin J. Evans, and Oliver Smithies performed groundbreaking research on homologous recombination in mouse stem cells, and they shared the 2007 Nobel Prize in Physiology or Medicine for their findings.<sup id="cite_ref-Nobel_2007_5-0" class="reference"><a href="#cite_note-Nobel_2007-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Traditionally, <a href="/wiki/Homologous_recombination" title="Homologous recombination">homologous recombination</a> was the main method for causing a gene knockout. This method involves creating a <a href="/wiki/DNA_construct" title="DNA construct">DNA construct</a> containing the desired mutation. For knockout purposes, this typically involves a drug resistance marker in place of the desired knockout gene.<sup id="cite_ref-Hall_6-0" class="reference"><a href="#cite_note-Hall-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The construct will also contain a minimum of 2kb of <a href="/wiki/Sequence_homology" title="Sequence homology">homology</a> to the target sequence. The construct can be delivered to <a href="/wiki/Stem_cell" title="Stem cell">stem cells</a> either through <a href="/wiki/Microinjection" title="Microinjection">microinjection</a> or <a href="/wiki/Electroporation" title="Electroporation">electroporation</a>. This method then relies on the cell's own repair mechanisms to recombine the DNA construct into the existing DNA. This results in the sequence of the gene being altered, and most cases the gene will be <a href="/wiki/Translation_(genetics)" class="mw-redirect" title="Translation (genetics)">translated</a> into a nonfunctional <a href="/wiki/Protein" title="Protein">protein</a>, if it is translated at all. However, this is an inefficient process, as homologous recombination accounts for only 10<sup>−2</sup> to 10<sup>−3</sup> of DNA integrations.<sup id="cite_ref-Hall_6-1" class="reference"><a href="#cite_note-Hall-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-:1_7-0" class="reference"><a href="#cite_note-:1-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Often, the drug selection marker on the construct is used to select for cells in which the recombination event has occurred. </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Physcomitrella_knockout_mutants.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Physcomitrella_knockout_mutants.JPG/220px-Physcomitrella_knockout_mutants.JPG" decoding="async" width="220" height="109" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Physcomitrella_knockout_mutants.JPG/330px-Physcomitrella_knockout_mutants.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Physcomitrella_knockout_mutants.JPG/440px-Physcomitrella_knockout_mutants.JPG 2x" data-file-width="2816" data-file-height="1391" /></a><figcaption>Wild-type <a href="/wiki/Physcomitrella_patens" title="Physcomitrella patens"><i>Physcomitrella</i></a> and <a href="/wiki/Knockout_moss" title="Knockout moss">knockout mosses</a>: Deviating <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a> induced in gene-disruption library transformants. <i>Physcomitrella</i> wild-type and transformed plants were grown on minimal Knop medium to induce differentiation and development of <a href="/wiki/Gametophore" title="Gametophore">gametophores</a>. For each plant, an overview (upper row; scale bar corresponds to 1 mm) and a close-up (bottom row; scale bar equals 0.5 mm) are shown. A: Haploid wild-type moss plant completely covered with leafy gametophores and close-up of wild-type leaf. B–E: Different mutants.<sup id="cite_ref-Egener_2002_8-0" class="reference"><a href="#cite_note-Egener_2002-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup></figcaption></figure> <p>These stem cells now lacking the gene could be used <a href="/wiki/In_vivo" title="In vivo">in vivo</a>, for instance in mice, by inserting them into early embryos. If the resulting chimeric mouse contained the genetic change in their germline, this could then be passed on offspring.<sup id="cite_ref-Hall_6-2" class="reference"><a href="#cite_note-Hall-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>In <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a> organisms, which contain two <a href="/wiki/Allele" title="Allele">alleles</a> for most genes, and may as well contain several related genes that collaborate in the same role, additional rounds of transformation and selection are performed until every targeted gene is knocked out. <a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a> may be required to produce <a href="/wiki/Homozygous" class="mw-redirect" title="Homozygous">homozygous</a> knockout animals. </p> <div class="mw-heading mw-heading3"><h3 id="Site-specific_nucleases">Site-specific nucleases</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=5" title="Edit section: Site-specific nucleases"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Frameshift_mutations_(13080927393).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/01/Frameshift_mutations_%2813080927393%29.jpg/303px-Frameshift_mutations_%2813080927393%29.jpg" decoding="async" width="303" height="214" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/01/Frameshift_mutations_%2813080927393%29.jpg/455px-Frameshift_mutations_%2813080927393%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/01/Frameshift_mutations_%2813080927393%29.jpg/606px-Frameshift_mutations_%2813080927393%29.jpg 2x" data-file-width="842" data-file-height="595" /></a><figcaption>Frameshift mutation resulting from a single base pair deletion, causing altered amino acid sequence and premature stop codon</figcaption></figure><p>There are currently three methods in use that involve precisely targeting a DNA sequence in order to introduce a double-stranded break. Once this occurs, the cell's repair mechanisms will attempt to repair this double stranded break, often through <a href="/wiki/Non-homologous_end_joining" title="Non-homologous end joining">non-homologous end joining</a> (NHEJ), which involves directly ligating the two cut ends together.<sup id="cite_ref-:1_7-1" class="reference"><a href="#cite_note-:1-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> This may be done imperfectly, therefore sometimes causing insertions or deletions of base pairs, which cause <a href="/wiki/Frameshift_mutation" title="Frameshift mutation">frameshift mutations</a>. These mutations can render the gene in which they occur nonfunctional, thus creating a knockout of that gene. This process is more efficient than homologous recombination, and therefore can be more easily used to create biallelic knockouts.<sup id="cite_ref-:1_7-2" class="reference"><a href="#cite_note-:1-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p><div class="mw-heading mw-heading4"><h4 id="Zinc-fingers">Zinc-fingers</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=6" title="Edit section: Zinc-fingers"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Zinc-finger_nuclease" title="Zinc-finger nuclease">Zinc-finger nuclease</a></div> <p>Zinc-finger nucleases consist of DNA binding domains that can precisely target a DNA sequence.<sup id="cite_ref-:1_7-3" class="reference"><a href="#cite_note-:1-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Each zinc-finger can recognize codons of a desired DNA sequence, and therefore can be modularly assembled to bind to a particular sequence.<sup id="cite_ref-:2_9-0" class="reference"><a href="#cite_note-:2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> These binding domains are coupled with a <a href="/wiki/Restriction_enzyme" title="Restriction enzyme">restriction endonuclease</a> that can cause a double stranded break (DSB) in the DNA.<sup id="cite_ref-:1_7-4" class="reference"><a href="#cite_note-:1-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Repair processes may introduce mutations that destroy functionality of the gene.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (December 2023)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading4"><h4 id="TALENS">TALENS</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=7" title="Edit section: TALENS"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Transcription_activator-like_effector_nuclease" title="Transcription activator-like effector nuclease">Transcription activator-like effector nuclease</a></div> <p>Transcription activator-like effector nucleases (<a href="/wiki/TALENs" class="mw-redirect" title="TALENs">TALENs</a>) also contain a DNA binding domain and a nuclease that can cleave DNA.<sup id="cite_ref-:3_10-0" class="reference"><a href="#cite_note-:3-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The DNA binding region consists of amino acid repeats that each recognize a single base pair of the desired targeted DNA sequence.<sup id="cite_ref-:2_9-1" class="reference"><a href="#cite_note-:2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> If this cleavage is targeted to a gene coding region, and NHEJ-mediated repair introduces insertions and deletions, a frameshift mutation often results, thus disrupting function of the gene.<sup id="cite_ref-:3_10-1" class="reference"><a href="#cite_note-:3-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="CRISPR/Cas9"><span id="CRISPR.2FCas9"></span>CRISPR/Cas9</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=8" title="Edit section: CRISPR/Cas9"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/CRISPR" title="CRISPR">CRISPR</a></div> <p>CRISPR (Clustered Regularly Interspaced Short Palindromic Repeats) is a genetic engineering technique that allows for precise editing of the genome. One application of CRISPR is gene knockout, which involves disabling or "knocking out" a specific gene in an organism.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (December 2023)">citation needed</span></a></i>]</sup> </p><p>The process of gene knockout with CRISPR involves three main steps: designing a guide RNA (gRNA) that targets a specific location in the genome, delivering the gRNA and a Cas9 enzyme (which acts as a molecular scissors) to the target cell, and then allowing the cell to repair the cut in the DNA. When the cell repairs the cut, it can either join the cut ends back together, resulting in a non-functional gene, or introduce a mutation that disrupts the gene's function. </p><p>This technique can be used in a variety of organisms, including bacteria, yeast, plants, and animals, and it allows scientists to study the function of specific genes by observing the effects of their absence. CRISPR-based gene knockout is a powerful tool for understanding the genetic basis of disease and for developing new therapies. </p><p>It is important to note that CRISPR-based gene knockout, like any genetic engineering technique, has the potential to produce unintended or harmful effects on the organism, so it should be used with caution.<sup id="cite_ref-:2_9-2" class="reference"><a href="#cite_note-:2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><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> The coupled Cas9 will cause a double stranded break in the DNA.<sup id="cite_ref-:2_9-3" class="reference"><a href="#cite_note-:2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Following the same principle as zinc-fingers and TALENs, the attempts to repair these double stranded breaks often result in frameshift mutations that result in an nonfunctional gene.<sup id="cite_ref-:2_9-4" class="reference"><a href="#cite_note-:2-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Non invasive CRISPR-Cas9 technology has successfully knocked out a gene associated in depression and anxiety in mice, being the first successful delivery passing through the <a href="/wiki/Blood%E2%80%93brain_barrier" title="Blood–brain barrier">blood–brain barrier</a> to enable gene modification.<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> </p> <div class="mw-heading mw-heading3"><h3 id="Knock-in">Knock-in</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=9" title="Edit section: Knock-in"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Gene_knock-in" title="Gene knock-in">Gene knock-in</a></div> <p>Gene knock-in is similar to gene knockout, but it replaces a gene with another instead of deleting it.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (December 2023)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=10" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Conditional_knockouts">Conditional knockouts</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=11" title="Edit section: Conditional knockouts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Conditional_gene_knockout" title="Conditional gene knockout">Conditional gene knockout</a></div> <p>A conditional gene knockout allows gene deletion in a tissue in a tissue specific manner. This is required in place of a gene knockout if the null mutation would lead to <a href="/wiki/Embryonic_death" class="mw-redirect" title="Embryonic death">embryonic death</a>,<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> or a specific tissue or cell type is of specific interest. This is done by introducing short sequences called loxP sites around the gene. These sequences will be introduced into the germ-line via the same mechanism as a knockout. This germ-line can then be crossed to another germline containing <a href="/wiki/Cre_recombinase" title="Cre recombinase">Cre-recombinase</a> which is a viral enzyme that can recognize these sequences, recombines them and deletes the gene flanked by these sites.<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> Other recombinases have since been created and employed in conditional knockout experiments.<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> </p> <div class="mw-heading mw-heading2"><h2 id="Use">Use</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=12" title="Edit section: Use"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Knockoutmouse80-72.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/Knockoutmouse80-72.jpg/220px-Knockoutmouse80-72.jpg" decoding="async" width="220" height="114" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/34/Knockoutmouse80-72.jpg/330px-Knockoutmouse80-72.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/34/Knockoutmouse80-72.jpg/440px-Knockoutmouse80-72.jpg 2x" data-file-width="772" data-file-height="401" /></a><figcaption>A <a href="/wiki/Knockout_mouse" title="Knockout mouse">knockout mouse</a> (left) that is a model of obesity, compared with a normal mouse</figcaption></figure> <p>Knockouts are primarily used to understand the role of a specific <a href="/wiki/Gene" title="Gene">gene</a> or <a href="/wiki/DNA" title="DNA">DNA</a> region by comparing the knockout <a href="/wiki/Organism" title="Organism">organism</a> to a <a href="/wiki/Wildtype" class="mw-redirect" title="Wildtype">wildtype</a> with a similar <a href="/wiki/Heredity" title="Heredity">genetic</a> background.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (December 2023)">citation needed</span></a></i>]</sup> </p><p>Knockout <a href="/wiki/Organisms" class="mw-redirect" title="Organisms">organisms</a> are also used as <a href="/wiki/Screening_(medicine)" title="Screening (medicine)">screening</a> tools in the development of <a href="/wiki/Drugs" class="mw-redirect" title="Drugs">drugs</a>, to target specific <a href="/wiki/Biological_processes" class="mw-redirect" title="Biological processes">biological processes</a> or <a href="/wiki/Deficiency_(medicine)" title="Deficiency (medicine)">deficiencies</a> by using a specific knockout, or to understand the <a href="/wiki/Mechanism_of_action" title="Mechanism of action">mechanism of action</a> of a <a href="/wiki/Drug" title="Drug">drug</a> by using a <a href="/wiki/Library" title="Library">library</a> of knockout <a href="/wiki/Organisms" class="mw-redirect" title="Organisms">organisms</a> spanning the entire <a href="/wiki/Genome" title="Genome">genome</a>, such as in <i><a href="/wiki/Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i>.<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> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=13" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Essential_gene" title="Essential gene">Essential gene</a></li> <li><a href="/wiki/Gene_knockdown" title="Gene knockdown">Gene knockdown</a></li> <li><a href="/wiki/Conditional_gene_knockout" title="Conditional gene knockout">Conditional gene knockout</a></li> <li><a href="/wiki/Germline" title="Germline">Germline</a></li> <li><a href="/wiki/Gene_silencing" title="Gene silencing">Gene silencing</a></li> <li><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a></li> <li><a href="/wiki/Planned_extinction" class="mw-redirect" title="Planned extinction">Planned extinction</a></li> <li><a href="/wiki/Recombineering" title="Recombineering">Recombineering</a></li> <li><a href="/wiki/Myostatin" title="Myostatin">Myostatin</a></li> <li><a href="/wiki/Belgian_Blue" title="Belgian Blue">Belgian Blue</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Gene_knockout&action=edit&section=14" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style 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(June 2004). <a rel="nofollow" class="external text" href="https://academic.oup.com/nutritionreviews/article-lookup/doi/10.1111/j.1753-4887.2004.tb00046.x">"Methods in Nutrition Science: Cre/loxP System for Generating Tissue-specific Knockout Mouse Models"</a>. <i>Nutrition Reviews</i>. <b>62</b> (6): 243–246. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1753-4887.2004.tb00046.x">10.1111/j.1753-4887.2004.tb00046.x</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nutrition+Reviews&rft.atitle=Methods+in+Nutrition+Science%3A+Cre%2FloxP+System+for+Generating+Tissue-specific+Knockout+Mouse+Models&rft.volume=62&rft.issue=6&rft.pages=243-246&rft.date=2004-06&rft_id=info%3Adoi%2F10.1111%2Fj.1753-4887.2004.tb00046.x&rft.aulast=Kos&rft.aufirst=Claudine+H.&rft_id=https%3A%2F%2Facademic.oup.com%2Fnutritionreviews%2Farticle-lookup%2Fdoi%2F10.1111%2Fj.1753-4887.2004.tb00046.x&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene+knockout" class="Z3988"></span></span> </li> <li id="cite_note-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-15">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTianZhou2021" class="citation journal cs1">Tian, Xueying; Zhou, Bin (January 2021). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8050033">"Strategies for site-specific recombination with high efficiency and precise spatiotemporal resolution"</a>. <i>Journal of Biological Chemistry</i>. <b>296</b>: 100509. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jbc.2021.100509">10.1016/j.jbc.2021.100509</a></span>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0021-9258">0021-9258</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8050033">8050033</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/33676891">33676891</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Biological+Chemistry&rft.atitle=Strategies+for+site-specific+recombination+with+high+efficiency+and+precise+spatiotemporal+resolution&rft.volume=296&rft.pages=100509&rft.date=2021-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8050033%23id-name%3DPMC&rft.issn=0021-9258&rft_id=info%3Apmid%2F33676891&rft_id=info%3Adoi%2F10.1016%2Fj.jbc.2021.100509&rft.aulast=Tian&rft.aufirst=Xueying&rft.au=Zhou%2C+Bin&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8050033&rfr_id=info%3Asid%2Fen.wikipedia.org%3AGene+knockout" class="Z3988"></span></span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20120929010716/http://www-sequence.stanford.edu/group/yeast_deletion_project/deletions3.html">"YeastDeletionWebPages"</a>. 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href="https://web.archive.org/web/20111026040816/http://www.bioscience.org/knockout/knochome.htm">Frontiers in Bioscience Gene Knockout Database (available on archive only)</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20131015201902/http://www.knockoutmouse.org/">International Knockout Mouse Consortium</a></li> <li><a rel="nofollow" class="external text" href="https://www.komp.org">KOMP Repository</a></li> <li><a rel="nofollow" class="external autonumber" href="https://pubmed.ncbi.nlm.nih.gov/2548993/">[1]</a></li> <li><a rel="nofollow" class="external autonumber" href="https://pubmed.ncbi.nlm.nih.gov/27798100/">[2]</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output 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class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_crops" title="Genetically modified crops">Crops</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_maize" title="Genetically modified maize">Maize/corn</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/MON_810" title="MON 810">MON 810</a></li> <li><a href="/wiki/MON_863" title="MON 863">MON 863</a></li> <li><a href="/wiki/Starlink_corn_recall#StarLink_corn" class="mw-redirect" title="Starlink corn recall">StarLink</a></li> <li><a href="/wiki/List_of_varieties_of_genetically_modified_maize" title="List of varieties of genetically modified maize">List of varieties of genetically modified maize/corn</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_potato" title="Genetically modified potato">Potato</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Amflora" title="Amflora">Amflora</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_rice" title="Genetically modified rice">Rice</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Golden_rice" title="Golden rice">Golden rice</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_soybean" title="Genetically modified soybean">Soybean</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Roundup_ready_soybean" class="mw-redirect" title="Roundup ready soybean">Roundup ready soybean</a></li> <li><a href="/wiki/Vistive_Gold" title="Vistive Gold">Vistive Gold</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_tomato" title="Genetically modified tomato">Tomato</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Flavr_Savr" title="Flavr Savr">Flavr Savr</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cotton#Genetic_modification" title="Cotton">Cotton</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bt_cotton" title="Bt cotton">Bt cotton</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_wheat" title="Genetically modified wheat">Wheat</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/HB4_Wheat" title="HB4 Wheat">HB4</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Arctic_Apples" title="Arctic Apples">Apple</a></li> <li><a href="/wiki/Arabidopsis_thaliana" title="Arabidopsis thaliana">Arabidopsis</a></li> <li><a href="/wiki/Genetically_modified_brinjal" title="Genetically modified brinjal">Brinjal</a></li> <li><a href="/wiki/Genetically_modified_canola" title="Genetically modified canola">Canola</a></li> <li><a href="/wiki/Papaya#Genetically_engineered_cultivars" title="Papaya">Papaya</a></li> <li><a href="/wiki/Blue_rose" title="Blue rose">Rose</a></li> <li><a href="/wiki/SmartStax" title="SmartStax">SmartStax</a></li> <li><a href="/wiki/Genetically_modified_sugar_beet" title="Genetically modified sugar beet">Sugar beet</a></li> <li><a href="/wiki/Tobacco#Genetic_modification" title="Tobacco">Tobacco</a></li> <li><a href="/wiki/Genetically_modified_tree" title="Genetically modified tree">Trees</a></li> <li><a href="/wiki/DMH-11_Mustard" title="DMH-11 Mustard">Mustard</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_animal" title="Genetically modified animal">Animals</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_mammal" title="Genetically modified mammal">Mammals</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetically_modified_mouse" title="Genetically modified mouse">Mouse</a> <ul><li><a href="/wiki/Knockout_mouse" title="Knockout mouse">Knockout mouse</a></li> <li><a href="/wiki/Oncomouse" title="Oncomouse">Oncomouse</a></li></ul></li> <li><a href="/wiki/Enviropig" class="mw-redirect" title="Enviropig">Enviropig</a></li> <li><a href="/wiki/Herman_the_Bull" class="mw-redirect" title="Herman the Bull">Herman the Bull</a></li> <li><a href="/wiki/Knockout_rat" title="Knockout rat">Knockout rat</a></li> <li><a href="/wiki/Alba_(rabbit)" title="Alba (rabbit)">Rabbit</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other animals</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetically_modified_insect" title="Genetically modified insect">Insects</a></li> <li><a href="/wiki/Genetically_modified_fish" title="Genetically modified fish">Fish</a> <ul><li><a href="/wiki/GloFish" title="GloFish">GloFish</a></li> <li><a href="/wiki/Genetically_modified_salmon" class="mw-redirect" title="Genetically modified salmon">Salmon</a></li></ul></li> <li><a href="/wiki/Genetically_modified_bird" class="mw-redirect" title="Genetically modified bird">Birds</a></li> <li><a href="/wiki/Genetically_modified_organism#Frogs" title="Genetically modified organism">Frogs</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetically_modified_bacteria" title="Genetically modified bacteria">Bacteria</a><br /> and <a href="/wiki/Genetically_modified_virus" title="Genetically modified virus">viruses</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ice-minus_bacteria" title="Ice-minus bacteria">Ice-minus bacteria</a></li> <li><a href="/wiki/Hepatitis_B_vaccine" title="Hepatitis B vaccine">Hepatitis B vaccine</a></li> <li><a href="/wiki/Oncolytic_virus" title="Oncolytic virus">Oncolytic virus</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Genetic_engineering_techniques" title="Genetic engineering techniques">Processes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">Inserting DNA</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Agrobacterium" title="Agrobacterium">Agrobacteria</a></li> <li><a href="/wiki/Gene_gun" title="Gene gun">Biolistics</a></li> <li><a href="/wiki/Electroporation" title="Electroporation">Electroporation</a></li> <li><a href="/wiki/Genetic_transduction" class="mw-redirect" title="Genetic transduction">Genetic transduction</a></li> <li><a href="/wiki/Lipofection" class="mw-redirect" title="Lipofection">Lipofection</a></li> <li><a href="/wiki/Microinjection" title="Microinjection">Microinjection</a></li> <li><a href="/wiki/Somatic_cell_nuclear_transfer" title="Somatic cell nuclear transfer">Somatic cell nuclear transfer</a></li> <li><a href="/wiki/Transfection" title="Transfection">Transfection</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Recombinant_DNA" title="Recombinant DNA">Recombinant DNA</a></li> <li><a href="/wiki/Transgenesis" class="mw-redirect" title="Transgenesis">Transgenesis</a></li> <li><a href="/wiki/Cisgenesis" title="Cisgenesis">Cisgenesis</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Uses</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">In agriculture</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetically_modified_food" title="Genetically modified food">Genetically modified food</a> <ul><li><a href="/wiki/Genetically_modified_food_controversies" title="Genetically modified food controversies">Controversies</a></li></ul></li> <li><a href="/wiki/Pharming_(genetics)" title="Pharming (genetics)">Pharming</a></li> <li>Companies <ul><li><a href="/wiki/BASF_Plant_Science" title="BASF Plant Science">BASF</a></li> <li><a href="/wiki/Bayer#Bayer_CropScience" title="Bayer">Bayer</a></li> <li><a href="/wiki/Dow_AgroSciences" title="Dow AgroSciences">Dow AgroSciences</a></li> <li><a href="/wiki/DuPont_Pioneer" class="mw-redirect" title="DuPont Pioneer">DuPont Pioneer</a></li> <li><a href="/wiki/Monsanto" title="Monsanto">Monsanto</a></li> <li><a href="/wiki/Syngenta" title="Syngenta">Syngenta</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">In <a href="/wiki/Human_genetic_engineering" class="mw-redirect" title="Human genetic engineering">humans</a> and<br /> diagnostics</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Gene_therapy" title="Gene therapy">Gene therapy</a></li> <li><a href="/wiki/Genetic_enhancement" class="mw-redirect" title="Genetic enhancement">Genetic enhancement</a></li> <li><a href="/wiki/Genetic_testing" title="Genetic testing">Genetic testing</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">In research</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a class="mw-selflink selflink">Gene knockout</a></li> <li><a href="/wiki/Gene_knockdown" title="Gene knockdown">Gene knockdown</a></li> <li><a href="/wiki/Gene_targeting" title="Gene targeting">Gene targeting</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related<br /> articles</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transgene" title="Transgene">Transgene</a></li> <li><a href="/wiki/Detection_of_genetically_modified_organisms" title="Detection of genetically modified organisms">Detection of genetically modified organisms</a></li> <li><a href="/wiki/Genetic_pollution" title="Genetic pollution">Genetic pollution</a></li> <li><a href="/wiki/Genetics_in_fiction" title="Genetics in fiction">Genetics in fiction</a></li> <li><a href="/wiki/Human_enhancement" title="Human enhancement">Human enhancement</a></li> <li><a href="/wiki/Reverse_transfection" title="Reverse transfection">Reverse transfection</a></li> <li><a href="/wiki/Transhumanism" title="Transhumanism">Transhumanism</a></li> <li><a href="/wiki/Genetic_use_restriction_technology" title="Genetic use restriction technology">Genetic use restriction technology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Regulation_of_genetic_engineering" title="Regulation of genetic engineering">Regulation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Cartagena_Protocol_on_Biosafety" title="Cartagena Protocol on Biosafety">Cartagena Protocol on Biosafety</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Geography</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Genetic_engineering_in_Europe" class="mw-redirect" title="Genetic engineering in Europe">Europe</a></li> <li><a href="/wiki/Genetic_engineering_in_Africa" class="mw-redirect" title="Genetic engineering in Africa">Africa</a></li> <li><a href="/wiki/Genetic_engineering_in_Asia" class="mw-redirect" title="Genetic engineering in Asia">Asia</a></li> <li><a href="/wiki/Genetic_engineering_in_North_America" class="mw-redirect" title="Genetic engineering in North America">North America</a> (<a href="/wiki/Genetic_engineering_in_the_United_States" class="mw-redirect" title="Genetic engineering in the United States">US</a>)</li> <li><a href="/wiki/Genetic_engineering_in_South_America" class="mw-redirect" title="Genetic engineering in South America">South America</a></li> <li><a href="/wiki/Genetic_engineering_in_Oceania" class="mw-redirect" title="Genetic engineering in Oceania">Oceania</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Similar fields</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Eugenics" title="Eugenics">Eugenics</a></li> <li><a href="/wiki/Cloning" title="Cloning">Cloning</a></li> <li><a href="/wiki/Stem_cell" title="Stem cell">Stem cell research</a></li> <li><a href="/wiki/Synthetic_biology" title="Synthetic biology">Synthetic biology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Biology" title="Biology">Biology</a></li> <li><a href="/wiki/Genetics" title="Genetics">Genetics</a></li> <li><a href="/wiki/Biotechnology" title="Biotechnology">Biotechnology</a></li> <li><a href="/wiki/Bioethics" title="Bioethics">Bioethics</a></li></ul> </div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Gene_knockout&oldid=1258539198">https://en.wikipedia.org/w/index.php?title=Gene_knockout&oldid=1258539198</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div 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